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IT IS a commonplace that novelty exercises such an attraction that it frequently 
diverts to itself a measure of attention out of all proportion to the true value of the 
subject or object. In science the field of every new discovery forthwith becomes 
the focal point round which attention centres, to the detriment of other fields more 
important but less glamorous. The tide of geographical exploration in the nineteenth 
century with its accompanying flood of zoological novelties exercised precisely this 
effect with the result that, whereas the vertebrate faunas of the Ethiopian, Oriental, 
Nearctic, and e\'en the Australian and Neotropical regions, have been more or less 
comprehensively listed in recent years, there have been few comparable works 
relating to the Palaearctic region where taxonomic zoology was born and cradled. 
The present work, whose geographical limits have been selected to link up with 
Chasen's (1940) list of Malayan mammals and Allen's (1939) similar list for the 
Ethiopian region, is an attempt to remedy this lack of balance in the field of 
systematic mammalogy. 

The authors have succeeded in producing a list which is not merely one of the 
working tools that every systematist must make for his own use. It is, in fact, a critical 
revision, shorn of all detailed argument, based on the unrivalled collections of the 

British Museum (Natural History) H. W. Parker 

London Keeper of ^oology 


In the course of reprinting the opportunity has been taken to incorporate the 
amendments contained in the sheet originally issued with the work, and those 
published later \nj. Mammal., 1953, 34: 516-518, as well as some which have since 
come to light. These amendments have, where possible, been made to the text itself; 
but those which were too long to be dealt with in this way have been printed as 
additional page 742d. 

A list of names which were overlooked in the original edition appears as additional 
pages 742a-c. Mr J. E. Hill has been responsible for this, as well as for in- 
corporating the amendments referred to above. 

No taxonomic alterations have been made, and it has not been possible in a 
piiotolithographic edition to deal with forms described since 1946. 

June 1 96-, 

British Museum (Natural Histoi"y) T. C. S. Morrison-Scott 

London Director 

Dedicated to the memory oj 



OUR late friend and colleague, James Lawrence Chaworth-Musters, had spent 
I much time latterly on the synonymies of the species of Palaearctic mammals, 
and in particular had devoted much patient research to the type localities 
and dates of publication of species described in the eighteenth and early nineteenth 
centuries. At the time of his death, in April 1948, he had nearly completed this work 
for the Insectivora and done much of the Chiroptera and Rodentia. His executors 
kindly placed his manuscript cards and foolscap sheets at our disposal, and we have 
made free use of the data referred to above. His death was a most untimely and un- 
fortunate loss to the Museum and to his friends and colleagues. (An obituary notice 
appears in Journal of Mammalogy, 1949, 30: 95.) 

Extent .\nd Method of this \Vork 

The area covered by this work is the Palaearctic region and the Indian^ and Indo- 
Chinese subdivisions of the Oriental region. Zoologists will be well aware of the 
difficulty in delimiting these zoogeographical areas. However, for the purposes of a 
list such as this, some arbitrary limit must be set. In Africa we have drawn the 
boundary along the parallel of 20° N. which, owing to the barrier of the Sahara, does 
correspond reasonably well with the facts. The boundary in Malaya has, however, 
been drawn in a purely arbitrary manner along the parallel of 10° N. This line has 
been chosen because it is the northern limit of the area covered by Chasen, 1940, 
Handlist of Malaysian Mammals. 

The hmits in point of time are from 1758 to 1946. That is to say, we have en- 
deavoured to include all forms of recent mammals named from the tenth edition of 
Linnaeus up tiU the end of 1946, except that domestic animals, and wild mammals 
which have become extinct, have as a rule been omitted. 

No one man can, of course, be a connoisseur of more than a small part of the class 
Mammalia. Nevertheless, in writing this work we have thought it worth while 
attempting a revision rather than making a mere nominal compilation. ^Ve have 
therefore re-examined all relevant monographs and revisions, in so far as they are 
known to us, together with the extensive study collections of the British Museum, and 
this checklist represents the results. Whether readers agree with our views or not, we 
hope that the presentation of such a survey within the covers of one book will prove 

There has been a considerable reduction in the number of named forms regarded 
as valid, though we have only proceeded with this "lumping" to the extent that the 
evidence before us justified it; there is probably much more to be done, and sub- 
species have been arranged in order of priority for the convenience of subsequent 

'The term 'India' has been used throughout in its zoogeographical sense to include the modern 
India and Pakistan. 


\Ve have recognized 809 species of mammals in the Palaearctic and Indian regions 
as defined above. 

^Ve have endeavoured to indicate the diagnostic characters of each genus and 
species by reference to the appropriate works, and where they are non-existent we 
have provided keys. The distribution of each species has been approximately shown, 
though it should be remembered that the distributions of many mammals are im- 
perfectly known and that the ranges of many of the larger mammals are shrinking 
every year. 


There are workers who seem to take a delight in bedevilling zoology with esoteric 
changes of nomenclature, to the considerable irritation of their colleagues and the 
confusion of non-specialists. In fact, exasperation at their efforts leads many to 
wonder whether they have any scientific work to attend to. 

Perhaps this unhappy circumstance is due to the idea that the only way to attain 
stability in nomenclature is rigorously to apply the law of priority, and that the 
resulting confusion will in the end have been worth while. It is o*" course true that 
with the passage of time the likelihood of fresh discoveries of early names becomes 
less. But the point is that the risk can never be eliminated. 

On the other hand, the Official List of Generic Names in ^oology and the Official List of 
Specific Trivial Names in ^oology do offer a chance of real stability (without confusion), 
and it is the view of the International Commission on Zoological Nomenclature that 
this is the way to attain it (Bull. ^ool. NomencL, 1950, .^.- 267, 627 and 5; 147J. It 
should therefore be the purpose of zoologists to see that the names of as many genera 
and species as possible of the groups in which they specialize are placed on these lists 
by the International Commission, and thereby protected from the activities of 
nomenclatorial excavators. 

The corollary to the above lists are the Official Index of Rejected and Invalid Generic 
Names in ^oologv and the Offcial Index of Rtjected and Invalid Specific Trivial Names in 
Zoology which the Commission instituted for the reception of names which they have 
either suppressed under their plenary powers, or declared to be otherwise un- 
available {Bull. Zool. NomencL, 1950, 4: 333). 

The Commission have urged that zoologists who discover a name which would 
cause confusion or inconvenience, through antedating a later but currently adopted 
name, should refrain from publishing their unfortunate find, and instead should 
hurry it off to the Commission for burial in the appropriate Index, at the same time 
requesting the Commission to place on the appropriate List the later but currently 
used name [Bull. ^ool. NomencL, 1950, 4: 234, j.' 18). 

These are the principles which we have endeavoured to follow in this checklist. So 
far as Palaearctic and Indian genera are concerned, the following works have proved 
the most troublesome : 

■a] Fiiscli, 1775, Dai NalursTslem der vierfussigen Ttiiere.^ This work has generally 
been regarded as unavailable under the Regies and Sherborn rejected it when com- 
piling his Index Animalium. Simpson (1945), however, in his Classification of Mammals 

' Formally irirdcd bv ()|>iuii>n 258, 1954. Ne\ii lluUis l)v ( )|)iniu,i -,8i, igbo, tin- C;omniibsion 
validated 0<iina Iristli. lyy^, ioi the European Fallow Deer. 


dates some fifteen well-known names from Frisch (1775). It is not clear why he did 
this since, in any case, some of the names have been dated from other authors by 
Opinion 91 of the International Commission. The matter has now been settled by 
the Commission who, in Paris in July, 1948, declared this work of Frisch to be un- 
available [Bull. Zool- Momencl., 1950, 4: 549)- The Commission made one reservation 
They had previously (Bull. ^ool. NomencL, 1950, 4: 547) declared that Zimmermann, 
■777) Specimen ^oologiae Geographicae was unavailable and that Zimmermann, 1778- 
I 783, Geographische Geschichte was available. The result of all this is that the name 
Dama becomes the technically valid name for the Virginian Deer of America instead 
offer the Fallow Deer of Europe, in which latter sense it has been used for years. The 
Commission, realizing the confusion which this would cause, indicated [Bull. Z^ol. 
NomencL, 1950, 4: 551) that they would use their plenary powers to prevent such a 
transfer if zoologists so desired, and in the meantime recommended them to 
make no change. Apart from this one name, the non-availability of Frisch (1775) 
appears to cause no inconvenience. 

[b] Oken, 1815-1816, Lehrbuch der Xaturgeschichte. This work can scarcely be held 
consistently to exhibit the principles of binominal nomenclature and the Commission 
are considering the question of its availability. ■ If Oken is declared unavailable, then 
there are certain generic names which it appears important to us to save. One of us 
(T. C. S. M.-S.) has therefore made application to the Commission for the following 
names- of Oken to be placed on the Official List : 

Citellus Tayra 

Genetta Vulpes 

Grison Pan 


(c) Brisson, 1762, Regnum Animale. The genera proposed as new in this work have 
been generally accepted by mammalogists and are now well estabhshed. But the 
technical validity of the book under the Regies is doubtful and the matter is now'before 
the Commission [Bull. J^ool. NomencL, 1950, 4: 314). In the meantime Hopwood, 
1947, P.^.S. iiy: 533, has rejected Brisson (1762) and would date his names from 
other and later authors. However, his suggestions, if adopted, would in several cases 
prove unfortunate, and we have asked the Commission to validate the following of 
the generic names'' of Brisson : 

Cuniculus. This is the Paca. The ne.\t use o{ Cuniculus is of Gronovius (1763) which, 
though also the Paca, seems insecure under the Regies. The next use is Cuniculus, 
Meyer (1790), which is the European Rabbit. It seems desirable, therefore, to retain 
Cuniculus Brisson. 

Glis. Unless Glis Brisson is validated, the name of the Fat Dormouse must be 
Myoxus Zimmermann (1780). (See Ellerman, 1949, Ann. Mag. N.H. 2: 894, who took 
the precaution of designating Glis zemni as the type species oi Glis Erxleben, 1777, in 
order to forestall the transference of Glis to the marmots, a worse confusion which 
would otherwise ensue from any suppression oi Glis Brisson.) 

Meles. It would be wise to validate this name as of Brisson in view of the doubt 
which surrounds the use as of GeoflTroy (1767) and Storr (1780). 

1 Rejected by Opinion 417, 1956. - A re-submission was requested for all applications prior to 

October 1959 which were still outstanding {Bull. ^ool. Aomeml., 1963, 20: 81. This has now been done, 
but in respect oi Pan and Panthera only (T.C.S.\1.-S.. 19651. '.No decision has yet (1963) been 

made. ■• Except for Odohenus these require a further application [fide Secretary to the Commission, 




Odubems. After considerable shuffling of the names of the Walrus, zoologists have 
riiially settled down with Oilohemis.' If this is inxuilid then Rosmarus Brunnich, 1771, 
will have to be used. 

Tragulus. The consequence of sinking this name ot Hrisson would indeed be un- 
fortunate. Hopwood suggests that Traaulus may equally well be dated from Boddaert 
' 178)). But Tragiihis Boddaert has nothing to do with the Tragulidae. It is Moschus 
moschifertti. a member of the Cervidae. A change in the family name of the che\T0tains 
would then become necessary, to add to the confusion. 

Tardinradiis. The earliest name for the Loris seems to be Tardixraihn Boddaert, 1 785, 
which has hitherto been regarded as preoccupied by Tardigradus Brisson, 1762, a 
Sloth. Hence Loris E. Geoftroy, 1796, is in current use for the Loris. \i Tardigradus 
Brisson is in\alid then Tardigradus Boddaert must be used for the Loris, which brings 
with it a secondary confusion in that the name "Tardigrada" is a synonym of 

Giraffa, Hyaena, Hydrochoerus, Lutra, Tapirus. These names are all available, with 
the same meaning, from Briinnich, 1771, ^oologiae Fuiidamentn, though the name of 
the C:apybara is here spelt Hydrochaeris. It may therefore be questioned whether there 
is any need to \alidate the use of these names from Brisson (1762). However, the 
Commission may well take the view that these names would be better protected by 
being \alidated from the earlier date, apart from the consideration of sanctioning a 
long-established usage. 

Pliivpus. This name comes in the same category as the last five, since it can be 
dated from Erxleben 1 1777) without change of meaning. There has, however, been 
some slight doubt about the type species and it is considered safer to validate the 
name as of Brisson !I762). 

d) Rafmesque, 1815, Analyse de la Nature. This book contains many nomina nuda, 
some of which are currently used. So far as the area covered by the present work is 
concei ncd, wc consider that one of these names, Muntiacus,- should be placed on the 
Official Lnt. The Muntjak was known many years ago as C.ervulus Blainville, 181 6, but 
Muntiacus is now in current use and, although it cannot really be pleaded that con- 
fusion would result, it would not be a helpful step to revert now to Cerrulus. We have 
submitted this case to the Commission. 

'Andersen, 1908, Ann. Mag. N.H. i: 431, discusses the technical availability of 
Rafinesque's (1815) genera.) 

.■\bbrevi.\tion's and Symbols 

The abbreviations of the titles of certain periodicals have been reduced beyond 
those shown in the World List: 

P.Z.S. = Proc. Zool. Soc, London 

.\.H. in combination) = Xat. Hist. 

A question mark Ix-lnre an entry in a synonymy docs not mean that the date is 
d(jubtful but that the name concerned is not certainly a synonym. 

A question mark in parentheses before the specific trivial name of a nominal race 

4 ' \ ahilatcd tjv ( )|iinMiii (I17, 11137. - \'.ilidalri.l bv Opinion 4(10, iij-,7. 


indicates that the latter is probably a race of the species concerned but that there is 
some doubt. 

N.V. =^ Non vidimus (with reference to the original publication). 


We gladly record our gratitude to many of our colleagues in this Museum for their 
generous help with, and friendly interest in, this work. 

We should especially like to thank the following: Dr. F. C. Fraser, for his advice 
and assistance in dealing with the Cetacea; Mr. A. C. Townsend, for helping us with 
difficult textual and bibUographical problems; and Mr. R. W. Hayman, for much 
help with the Chiroptera. 

So far as possible every reference in this book has been checked with the original, 
and we desire to record the assistance which has been given us by the following of our 
colleagues — in fact without their help this work would almost certainly have proved 
too much for us: Mr. R. \V. Hayman and Mr. G. ^V. C. Holt of the Mammal Room, 
who between them checked most of the references; Mr. G. ^V. F. Claxton, Mr. F. C. 
Sawyer, Mr. W. H. Mabbott and Mr. J. E. Yateman of the General and Zoological 
Libraries, to whom an incomplete or distorted reference was a professional challenge 
which they rarely failed to meet; and Miss J. M. Ingles who has been of great 
personal assistance to us. 

Authorship and New Names 

We take joint responsibility for this book except for the classification of the 
Rodents and Lagomorphs, which is the work of J. R. E., and the Ungulates for which 
T. C. S. M.-S. is responsible. 

The new names contained in this work, a list of which appears on page 742, are 
proposed by us jointly irrespectively of the order to which they belong. 

J. R. Ellerman 


British Museum (Natural History) 
2,1st December, 1950 



There are very few works dealing extensively with the class Mammalia. The 
following are the most important: 

Gregory, W. K. 19 io. The orders of mammals. Bull. Amer. Mus. M.H. sy. 
Flower, W. H., & Lydekker, R. i8gi. An introduction to the study of mammals, living 

and extinct. London (A. & C. Black). 
Parker, T. J., & Haswell, \V. A. 1940. A textbook of zoology, 2, Chordata. (Revised 

by C. Forster Cooper.) London (Macmillan). 
Simpson, G. G. 1945. The principles of classification and a classification of mammals. 

Bull. Amer. Mus. M.H. 85. 
Weber, M. 1927-1928. Die Sdugetiere (2 vols). Jena (G. Fischer). 
Winge, H. 1923-1924. Pattedyr-Slaegter (3 vols). Copenhagen (H. Hagerup). (English 

translation by G. M. Allen and E. Deichmann, 1941-1942. Copenhagen (C. A. 


Simpson (1945) is the basic work on the classification of mammals. The mammals 
with which this checklist is concerned all belong to the infraclass Eutheria, which 
Simpson divides into four cohorts; 


Orders: Insectivora, Dermoptera, Chiroptera, Primates, Pholidota. 


Orders: Lagomorpha, Rodentia. 


Order: Cetacea. 

Superorder: Ferae 

Order: Carnivora (Suborders: Fissipedia, Pinnipedia). 
Superorder: Paenungulata 

Orders: Proboscidea, Hyracoidea, Sirenia. 
Superorder: Mesaxonia 

Order: Perissodactyla. 
Superorder: Paraxonia 

Order: Artiodactyla. 

We agree with Simpson in distinguishing the Mutica and the Glires, and follow 
the broad outline of his classification except that we retain the Pinnipedia as an 
order, and on account of the fact that his Ferungulata seem closely allied to his 
Unguiculata we have listed them directly after this cohort. 


ORDERS: i. Inscctivora, page 8 

2. Dermoptcra, page 89 

3. Chiroptera, page 90 

4. Primates, page 189 
-). Phi ilidota, page 213 

6. Clarnivora, page 215 

7. Pinnipedia, page 321 

8. Hyracoidea, page 334 

9. Proboscidea, page 336 
10. Sirenia, page 337 

I I . Perissodactyla, page 338 

12. Artiodactyla, page 343 

13. Lagomorpha, page 419 

14. Rodentia, page 456 

15. Cletacea, page 712 


Special works of rel'ercnce: Besides works such as G. S. Miller, 1912, Catalogue of 
the Mammals of Western Europe; G. M. Allen, 1938 & 1940, Mammals oj China and 
Moncolia; and works by Bobrinskii and Ognev on Mammals of the U.S.S.R., see 
particularly A. Cabrera, 1925, Genera Mammaltum; Insectivora, Galeopithecia. This work 
gives keys to all families and genera of Insectivora here recognized and dealt with. 
See also G. E. Dobson, 1 882-1 890, Monograph oJ the Insectivora. 

FAMILIES: Erinaceidae, page 16 

Macroscelididae, page 14 
Soricidae, page 41 
Talpidac, page 29 
Tupaiidae, page 9 

Simpson, 1945, Bull. Amer. Mus. N.H. 8j: 61, 176, 182, referred the Tupaiidae (as 
type of a special superfamily), to the suborder Prosimii of the order Primates. Most 
authors refer these animals to the Insectivora. If they are so close to Lemuroids that 
it is thought best to refer them to Primates, surely another course would be to refer 
the Prosimii to the Insectivora, and restrict Primates to the Anthropoidea (perhaps 
with the Tarsiidae). Some authors, such as Gregory and AVeber, separate the 
Tupaiidae and Macroscelididae from the Insectivora as a separate order Menotyphla. 
This is strongly supported by Broom {in litl). However, for the present wc preicr to 
list these families as Insectivora. Apart from Tupaiidae Simpson recognizxd three 
superfamihes : the Erinaceoidea for the Erinaceidae and some extinct allies; the 
Macroscelidoidea for the Macroscelididae (which only occur in North-VVest Africa in 
the present region) ; and the Soricoidea for the Soricidae and Talpidae (which appear 
to us to be very distinct from each other morphologically, particularly as regards the 
very large first lower incisor in the Soricidae). 



Genera: Anathana, page 13 
Dendrogale, page 13 
Tupaia, page 10 

This family was monographed in great detail by Lyon, 191 3, Proc. U.S. Nat. Mus. 
4§: 1-188. Most subsequent classifications have been based on this useful paper. Only 
the typical subfamily, the Tupaiinae, occurs within the region now under discussion, 
and its distribution is Indo- Malayan. Lyon gives keys to generic characters of the 
three genera listed above and their extralimital allies. The main distinctions of the 
four species here listed as valid and which are certainly known to occur north of the 
area treated by Chasen, 1 940, Handlist Malaysian Mammals, are as follows : 

1. Relatively small animals, with the tail rounded and close-haired for its whole 

length. Dendrogale munna 

Relatively larger animals, with the tail clothed with longer hairs, and squirrel-like 

in formation 2 

2. Lower canine little differentiated, not higher than adjacent lower I 3 and P 2. 

Fenestrae in zygoma small and inconspicuous; hypocones in upper molars un- 
usually prominent. Anathana ellioli 
Lower canine clearly differentiated, clearly higher than adjacent lower I 3 and 
P 2. Fenestrae in zygoma normally large and conspicuous; hypocones in upper 
molars most often less prominent. 3 

3. Tail considerably longer than head and body. Much black on lower part of back. 

Lower canine much larger than the incisor in front of it; central upper incisors 
conspicuously larger than lateral pair. Tupaia mcobarica 

Tail most often shorter than, or not much longer than, head and body. Colour of 
back different. Lower canine and central upper incisors not conspicuously 
enlarged. Tupaia glis 

(We have not included Tupaia minor in the key as we are not sure whether it is 
extralimital or not. According to Lyon's key, T. minor should be dentally as nicobarica 
but smaller than that species and coloured differently.) 

North of the Malay Peninsula Lyon recognized two species, T. glis and T. belangeri, 
in addition to the very distinct T. nicobarica. They were said to differ in colour and 
mammary formula. But since Lyon's revision was published there have been many 
new forms described of the T. glis group, and examination of the types in the British 
Museum alone shows that there is no certain colour distinction between belangeri and 
races referrable to glis. Chasen (1940) refers several of Lyon's species to T. glis as 
races, and it seems that there is little essential difference between the southern glis 
races and the northern belangeri and allies, which are here considered as representing 
T. glis. It may be noted that, with reference to the above key, the hypocones may be 
present in the upper molars of some individuals of T. glis siccata which in this character 
approaches Anathana; and that in some forms of T. glis, for instance T. g. lepcha, there 
is a tendency for the tail to be longer than the head and body. The retention of the 

palaearc:tk: and Indian mammals i7r,8-i946 

genus Atiathana is here principally based on the reduced lower canine. Thomas (191 7) 
thought two forms of the T. glis group occurred in Tenasserim. These two, clanssa 
and tenaster, differ in the length of the rostrum, which is more lengthened in 
However, these two forms look so alike externally that very tentatively tenaster is here 
regarded as a synonym. To prove the contrary it would be necessary to collect a 
much larger series in Tenasserim than these two names are based on. 

SuBF.^iMii.Y T u p a i i n a e 

Genus TUPAIA Raffles, 1821 

182 1. Tupaia Rafiles. Trans. Linn. Soc, London, 13: 256 (May, 182 1.) Tupaia 
fcrruguna Raffles. 

1821. Sorex-aih Cuvicr cS: Gcotfroy, Hist. Nat. Mamm. livraison, 35: i (Deicmbcr, 

1821, or perhaps early in 1822.) Snrex ofis Diard & Du\auccl. 

1822. Glisorex Desmarest, Mammalogie, footnote, 536. Substitute for Sorex-glis. 
1824. Cladobates Cuvier, Dents Mamm. 251, pi. 17. Tupaia ferruginea Raffles. 
1827. Hylogale Temminck, Mon. Mamm. xix. Substitute for Tupaia. 

1843. Hylogalea Miiller & Schlegel, \'erh. Nat. Gesch. Ned. Overz. Bezitt. 139. 

1855. Glisosorex Giche\, Odontographie, 18. (Emendation of G7;jorf.v.) 
i860. Tapaia Gray, Ann. Mag. N.H. 5: 71. (? Misprint for Tupaia.) 
1882. Gliriiorex Scudder, Nomencl. Zool. 2: 131. (Emendation of Glisorex.) 
1888. G///)ora Jentink, Cat. Syst. Mus. H.N. Pays Bas. 12, Mamm.: 118. Clipora 

leucogaster ]enX\nk {nom. nud.) = Tupaia minor Gunther. 
1913. Tana Lyon, Proc. U.S. Nat. Mus. 4;^: 134. Tupaia tana Raffles, from Sumatra. 
Valid as a subgenus. 
3 species in the area covered by this list: 
Tupaia glis, page i o 
Tupaia minor, page 1 2 
Tupaia nicoharica, page 12 

Tupaia glis Diard, 1820 Common Tree-Shrew 

Approximate distribution of species: Yunnan, Kwangsi, Hainan in South-West 
China; Sikkim, NLanipur, Assam, Burma, Tenasserim; Indo-China, Siam, .\Lalay 
States, Sumatra, Java, Borneo, and many small adjacent islands, to Palawan. 

(Tup.MA GLIS Gus Diard, 1820. Extralimitaj) 

1820. Sorex glis Diard, Asiat. J. Month. Reg. 10: 478. (N.V.,Jide Lyon cS: Chasen.) 

Penang Island, Malay Peninsula. 
1822. .So/r.v^/w Diard & Duvaucel, Asiatick Res. 14: 471, pi. o. Penang Island. 

Tupaia glis belangeri \Vagner, 1841 

1841. Cladobates belangeri Wagner, Schreber's Siiugeth. Suppl. 2: 42. Siriam, near 

Rangoon, Pegu, Burma. 

1842. Tupaia pegiianus Lesson, Nouv. Tabl. Regn. Anim. Mamm. 93. ? Pegu. 
Range: Southern Burma and certain islands of Mergui Archipelago. 



i860. Sciurus dissimilis Ellis in Gray, Ann. Mag. N.H. 5.- 71. Pulau Condore, off 
south coast of Indo-China. 

TuPAiA GLIS CHINENSIS Anderson, 1879 

1879. Tupaia chinensis Anderson, Zool. Res. West Yunnan, 129, pi. 7, figs. 8 and 9. 

Ponsee, Kakhyen Hills, 3,185 ft., and Muangla, Sanda Valley, 2,400 ft.. 

Western Yunnan, China. 

Tupaia glis modesta J. Allen, 1906 

1906. Tupaia modesta Allen, Bull. Amer. Mus. N.H. 22: 481. Lei-mui-mon, Island of 

Hainan, South China. 
1 9 14. Tupaia belangeri yunalis Thomas, Ann. Mag. N.H. ig: 244. Mongtsze (or 

Mengtsz), Southern Yunnan, China. (Status ^(/c Osgood, 1932.) 
1925. Tupaia belangeri tonquinia Thomas, P.Z.S. 497. Bao-ha, Tonkin, Indo-China. 

{Sta-tui fide Osgood, 1932.) 
(?) 1936. Tupaia belangeri pingi Ho, Contr. Biol. Lab. Sci. Soc. China, 12, 4: 78. 

Bao-peng, Island of Hainan. 
Range: Hainan, Annam, Laos, Tonkin, and Southern Yunnan. 

Tupaia glis concolor Bonhote, 1907 

1907. Tupaia concolor Bonhote, Abstr. P.Z.S. 2; P.Z.S. 7. Nhatrang, Annam, Indo- 

China. Ranges to Cambodia and Cochin-China. 

Tupaia glis siccata Thomas, 19 14 

1914. Tupaia belangeri siccata Thomas, Ann. Mag. N.H. ij: 243. Zibugaung, Lower 

Chindwin, Burma. Range includes Chin Hills, Mt. Popa,"Shan' States, 


Tupaia glis laotum Thomas, 19 14 

19 14. Tupaia belangeri laotum Thomas, Ann. Mag. N.H-. ij: 244. Nan, 290 m., Siam. 

Tupaia glis sinus Kloss, 1916 

1916. Tupaia concolor sinus Kloss, P.Z.S. 36. Koh Chang (Island), South-East Siam. 

Tupaia glis Clarissa Thomas, 191 7 

1917. Tupaia Clarissa Thomas, J. Bombay N.H. Soc. 2^: 200. Bankachon, Victoria 

Province, Tenasserim. 
(?) 1917. Tupaia belangeri tenaster Thomas, J. Bombay N.H. Soc. 25.- 201. Tagoot, 
Great Tenasserim River, Tenasserim. 

Tupaia glis cambodiana Kloss, 1919 

1919. Tupaia glis cambodiana Kloss, J. N.H. Soc. Siam, 3: 357. Klong Yai, South-East 

Tupaia glis olivacea Kloss, 1919 

1919. Tupaia glis olivacea Kloss, J. N.H. Soc. Siam, 3: 358. Pak Bu, near Tachin, 
Central Siam. 


1 92 1. Tupaia belangeri assamensis VVroughton, J. Bombay N.H. Soc. 2y: 599. Mokok- 

chung, 5,000 ft., Naga Hills, Assam. Range includes Manipur. 

Tup.A.i.'^ GLIS cocHiN'CHiNENSis Robinson & Kloss, 1922 

1922. Tupaia glis cochinchincnsis Robinson & Kloss, Ann. Mag. N.H. g: 87. Trangbom, 

30 miles cast of Saigon, Cochin-China. 


1922. Tupaia disiimilis annamensis Robinson & Kloss, Ann. Mag. N.H. g: 87. Daban, 
650 ft.. Southern Annam., Indo-China. 

Tupaia glis versurae Thomas, 1922 

1922. Tupaia belangeri versurae Thomas, J. Bombay N.H. Soc. 28: 42B. Dening, 
2,250 ft., Mishmi Hills, North Assam. 

Tupaia glis lepcha Thomas, 1922 

1922. Tupaia belangeri lepcha Thomas, J. Bombay N.H. Soc. 28: 428. Narbong, near 

Darjeeling, 2,000 ft. Ranges to Bhutan Duars. 

Tupaia glis bru.metta Thomas, 1923 

1923. Tupaia belangeri brunetta Thomas, J. Bombay N.H. Soc. 2g: 84. King Island, 

Mergui Archipelago. 

Tupaia nicobarica Zclebor, 1869 Nicobar Tree-Shrew 

Approximate distribution of species: Nicobar Islands, Bay of Bengal. 

Tup.MA nicobarica nicobarica Zelebor, 1869 

1869. Cladobates nicobaricus Zelebor, Reise Novara, Zool. Theii, /.• 17, pi. i, figs, i, 2, 
3, and pi. 2. Great Nicobar, Nicobar Islands. 

Tup,\iA nicobaric.\ surda Miller, 1902 

1902. Tupaia nicobarica surda Miller, Proc. U.S. Nat. Mus. 2^: 774. Little Nicobar, 
Nicobar Islands. 

Tupaia minor Giinther, 1876 Gunther's Tree-Shrew 

.Approximate distribution of species: Malay States, Sumatra, Borneo; north into 
South Siam. 

(Tupaia minor minor Gunther, 1876. Extralimital) 

1876. Tupaia w;«nr Giinther, P.Z.S. 426. Borneo, mainland opposite Island of Labuan. 

TuPAi.\ minor malaccana Anderson, 1879 

1879. Tupaia malaccana Anderson, Zool. Res. Yunnan, 134, pi. 7. Malacca. Chasen 
( 1940, 10) quotes two immature examples of T. minor from Koh Lak, South- 
\Vest Siam. We are unable to trace this locality, but have reason to believe 
it is just inside our region, and extralimital to the part of Peninsular Siam 
covered by Chasen. 


Status not sure : 

TuPAiA siAMENSis Gyldcnstolpe, 19 16 

1916. Tupaia siamensis Gyldcnstolpe, K. Svenska Vetensk. Akad. Handl. 57, 2; 20. 
Koh Lak, Siamese Malaya. From descriptions it is much like T. minor except 
for considerably larger size, but too small for nicobarica (head and body 
145 mm., tail 175 mm., hindfoot 42 mm.). There are no dental details in 
the original description. 

Genus ANATHANA Lyon, 1913 

19 1 3. Anathana Lyon, Proc. U.S. Nat. Mus. 4^: 120. Tupaia ellioti \Vaterhouse. 

I species: Anathana ellioti, page 13 

Lyon divided this genus into three nominal species, but we doubt whether they are 
really more than well differentiated races of the earliest named form. 

Anathana ellioti \Vaterhouse, 1850 Madras Tree-Shrew 

Approximate distribution of species: Eastern Ghats, Madras, Bihar, Central 
Provinces, Surat District, Bombay (part), in Peninsular India. 

Anathana ellioti ellioti Waterhouse, 1850 

1850. Titpaia ellioti Waterhouse, P.Z.S. i84g: 107, pi. Mamm. 13. Hills between 
Cuddapah and Nellore, Eastern Ghats, India. Range: Eastern Ghats and 
^ Shevaroy Hills, India. 

Anathana ellioti wroughtoni Lyon, 1913 

191 3. Anathana wroughtoni Lyon, Proc. U.S. Nat. Mus. ^5.- 123. Mandvi, near Bom- 
bay, India. Range: Region of Satpura Hills, and Dangs, near Bombay, 
Western India. 

Anathana ellioti pallida Lyon, 191 3 

1913. Anathana pallida Lyon, Proc. U.S. Nat. Mus. 4^: 124. Munbhum, Bihar, India. 

Range: Raipur in Central Provinces north-eastwards as far as the Ganges, 


Genus DENDROGALE Gray, 1848 

1848. Dendrogale Gray, P.Z.S. 23. Hylogalea murina Schlegel & Mtiller. 

I species in the area covered by this list: 

Dendrogale murina, page 14 

Lyon divided this genus into two groups. The murina group is characterized as 
having hght colour, face markings present, and small claws. D. murina was supposed 
to have come from Borneo, and the Indo-Chinese species is currently called D.frenata. 
However, Chasen, 1940, Handlist Malaysian Mammals, 10, states: ^'Dendrogale murina 
. . . said to have come from Pontianak, West Borneo, seems a very doubtful species 



(i.e., of doubtful occurrence in the Malaysian region); it has never again turned 
up in Borneo and the type is so very hke the Indo-Chinese frenala that I have dropped 
the name from the Malaysian list." Lyon (p. 131) suggests that there is "just a 
possibility that the type oi miirina is an example oi frenala wrongly labelled as coming 
from Pontianak, Borneo". He states that Dr. VV. L. Abbott, with much careful 
collection in the neighbourhood of the supposed (Bornean) type locality, failed to 
secure additional specimens ot miirina. It seems logical, therefore, to adopt the name 
miirina, which antedates y/rwa/rt by seventeen years, for the Indo-Chinese species. 

Dendrogale murina Schlcgel & MiiUer, 1845 Northern Smooth-tailed Trcc-Shrew 

Approximate distribution of species: Cambodia, Annam, Cochin-China. in Indo- 

Dendrogale murina Schlcgel & Mtiller, 1845 

1845. Hrlogalfa murina Schlcgel & Mtiller, Verh. Nat. Gesch. Ned. Overz. Bezitt. 

167, pi. 26, fig. 5; pi. 27, figs. 17-18. Supposed to be from Pontianak, West 

Borneo (error ?). 
(?) i860. Ttipaia frcnata Gray, Ann. Mag. N.H. 6: 217. Cambodia, Indo-China. 


Genus: F.hjiliantulus, page 15 

This family is principally from South and East Africa, but one of the species occurs 
in Morocco and Algeria. The genus differs from those tropical genera which ante- 
date it roughly as follows. In Rhynchocvon, which contains large species, the hallu.x is 
absent; the dentition is abnormal, in that the upper incisors are reduced to one, 
which is nearly vestigial, so that there are no functional front teeth in front of the 
canine, which is conspicuously enlarged and dominant. In the other genera there are 
three upper incisors and the upper canine is not extremely dominant. Petrodromus 
contains large species with no hallux and with normal dentition. Macroscelides con- 
tains small species with the hallux small but clawed and present, and the bullae 
enormously enlarged. Elephantulus is like Macroscelides but with quite normal, small 
bullae. Usually it has ten lower (and ten upper) cheekteeth, thereby differing from 
Nasilio which is closely allied but which has normally ele\en lower cheekteeth. 
Perhaps Nasilio is only a subgenus oi Elephantulus. 

The Palaearctic species oi Elephantulus is the first specific name in the genus. There 
arc three rather well-defined groups of species in British Museum material of 
Elephantulus. E. intuji from South Africa stands apart from all the remainder in having 
the upper P 3 (the fifth tooth from the back) large, four-cusped and molariform. For 
this, the subgeneric name Elephantomys Broom, 1937, is available. E. rupestris, South 
African, the type, has the upper P 3 narrow, sectorial, and usually two-cusped. Two 
co-types of E. rupestris are in the British Museum, and both show the characters 
clearly. The bullae in these are broken, but in other specimens, and in types of forms 
named as subspecies of rupestris, the bullae are somewhat llattened so that the 



external part of the bulla is about on the same level with the median part of the bulla 
as seen in ventral view. E. rupestris myurus and E. rupestris jamesoni are subspecies 
represented in London, and E. capensis belongs to the same group. The remainder 
have the upper P 3 narrow and sectorial but the bullae are not flattened, so that the 
e.xternal part of the bulla is on a much lower level than the median part of buUa 
as seen in ventral view. E. rozeti is the prior name for this group, and the following 
types have been available for examination: atlantis, clivorum, deserti and moralus. 
Essentially similar forms are represented by the types oi peasei and somalicus (respec- 
tively from Abyssinia and Somaliland), boramis, delicatus and dundasi (all from Kenya), 
ocularis, pulcher and renatus (all from Tanganyika). The type of edwardsii has P 3 as in 
rupestris, but the bullae are broken. All these forms are listed by G. Allen (1939). 

Genus ELEPHANTULUS Thomas & Schwann, 1906 

1906. Elephantulus Thomas & Schwann, Abstr. P.Z.S., No. 33, 10. P.Z.S. 577. 

Macroscelides rupestris Smith, from the Cape Province. 
1937. Elephaniomys Broom, S. Afr. J. Sci. jjj.- 758. E. langi Broom from cave deposits 

at Schurveberg, Transvaal. Valid as a subgenus, to include also the living 

species E. intufi Smith from the West Transvaal. 

I species in the area covered by this list : 
Elephantulus rozeti, page 15 

Elephantulus rozeti Duvernoy, 1833 North African Elephant-Shrew 

Approximate distribution of species : Morocco and Algeria. Closely allied forms 
(subspecies ?) inhabit Abyssinia, Somaliland, Kenya and Tanganyika. 

Elephantulus rozeti rozeti Duvernoy, 1833 

1833. Macroscelides rozeti Ttuwemoy , Mem. Soc. Hist. Nat. Strasbourg, /, 2: art. 4, 18, 
pis. I, 2. Near Oran, Algeria. Range: Northern .\lgeria, Oran, Northern Rif 

Elephantulus rozeti deserti Thomas, 1901 

1 90 1. Macroscelides rozeti deserti Thomas, Ann. Mag. N.H. 8: 155. Near Jebel Bourzel, 
Biskra, Algeria. 

Elephantulus rozeti atlantis Thomas, 1913 

1913. Elephantulus rozeti atlantis Thomas, Novit. Zool. 20: 587. Northern slope of 
Great Atlas of Morocco, south of Seskawa, Ain Moussa. 

Elephantulus rozeti moratus Thomas, 19 13 

1913. Elephantulus rozeti moratus Thomas, Novit. Zool. 20: 587. Jebel Chedar, about 

80 km. south-east of Mazagan, South- Western Morocco. Range includes 

desert of Zragna, Morocco. 

Elephantulus rozeti clivorum Thomas, 191 3 

1913. Elephantulus deserti clivorum Thomas, Novit. Zool. 20: 588. Guelt-es-Stel, 900 m., 

plateau of Eastern Algeria. Range: as above, also Matmata, Southern Tunis, 

and Maafa, Eastern Algeria. 




Genera : Echinoiorex, page 1 7 
Eriiiacfus, page 19 
HiDiiechinus, page 23 
Hylomys, page 17 
Meotelracus, page 18 
Paracchitius, page 26 

This family is divided into two subfamilies : the Echinosoricinae, containing Neo- 
tetracus, Echinosorex and Hylomys, and chiefly Indomalayan in distribution; and the 
Erinaceinae, containing the true Hedgehogs, Erinaceus, Paraechinus, Htmiechinus, which 
is principally Palaearctic and African in distribution. Formerly the Hedgehogs were 
all referred to a single genus Erinaceus Linnaeus, and this classification is still followed 
by some authors, for instance by Bobrinskii (1944). Thomas, 1918, Ann. Mag. 
N.H. i: 193-196, divided these animals into five genera. Of these we are not pre- 
pared to admit Atelcnx as more than a subgenus; and certainly not Aethechinus as 
anything but a synonym of Atelerix which was restricted by Thomas to species in 
which the small hallux is absent. This character is now known not to be constant: 
see J. A. Allen, 1922, Bull. Amer. Mus. N.H. 47: 13. But there seems a generic 
division in the formation of the bullae between Erinaceus and Herniechinus. From 
the last, Paraechinus is not so easily distinguished, and it might be regarded as 
only a subgenus of Herniechinus. The genera admitted here may be keyed as 
follows : 

1. Coat not definitely spiny; 10 or 1 1 lower teeth (40 or 44 teeth in all). Tail at least 

visible externally. (Subfamily Echinosoricinae) 2 

Coat densely spiny dorsally. Eight lower teeth (36 teeth in all). Tail not, or 
scarcely, apparent. (Subfamily Erinaceinae) 4 

2. Coat rough and harsh; much larger animal, head and body 265-345 mm. 

in B.M. material. Colour striking; typically mixed black and white, with 
black stripe round eye, and with tail dark basally, pale terminally. (White 
forms occur.) Tail long, averages over 80 per cent, of head and body. First 
upper incisor and upper canine strong and well dilferentiated. 

Coat soft; smaller animals, head and body in adults 143 mm. at most, and usually 
less, in the specimens examined. Colour drab; appearance very reminiscent of 
certain Voles (Microtinae). Tail short, averages 54 per cent., or less, of head and 
body in B.M. material. Upper canine weak (Hylomys) or scarcely differentiated 
(Neotetracus). 3 

3. Tail averages about 54 per cent, of head and body, and is usually over 60 mm. in 

length. Normally 10 upper and lower teeth. NEOTETRACUS 

Tail very short, averages 17 per cent, of head and body, and reaches 30 mm. in 
only one specimen of the material examined. Normally 1 1 upper and lower 
teeth. HYLOMrS 


4. "Pterygoids inflated, their cavity communicating with that of bullae. Paraptery- 

goid fossae shallow. Postglenoid fossae even larger and more hollowed out than 
Hemiechinus" (Thomas). PARAECHINUS 
Pterygoids and bullae more normal. 5 

5. Postglenoid process as large as mastoid process, hollow internally. 

Postglenoid process small, not hollowed out, and much surpassed by the mastoid 
process. ERINACEUS 

Subfamily Echinosoricinae 

Genus ECraNOSOREX Blainville, 1838 

1827. Gymnura Lesson, Man. Mamm. 171. Gymnura rafflesii Lesson = Viverra gymnura 
' Rafiles. Not of Kuhl, 1824. 

1838. Echino-sorex Blainville, C.R. Acad. Sci. Paris, 6: 742. Viverra gymnura Raffles. 
1840. Echinosorex VAa^nviWc, Osteogr., Insectiv. 109. Substitute for Echino-sorex. 

I species : Echinosorex gymnurus, page 1 7 

Echlnosorex gymnurus Raffles, 1821 Moonrat or Raffles' Gymnura 

Approximate distribution of species: Tenasserim, Lower Siam, Malay States, 
Sumatra, Borneo. 

(Echinosorex gymnurus gymnurus Raffles, 1821. Extralimital) 
182 1. Viverra gymnura Raffles, Trans. Linn. Soc. London, /j.- 272. Bencoolen, 
Sumatra. (Ranges to Malay Peninsula.) 

Echinosorex gymnurus birmanicus Trouessart, 1879 

1879. Gymnura birmanica Trouessart, Rev. ZooL Paris, 240. Bankachon, Southern 

1888. Gymnura raffiesi Blanford, Fauna Brit. India, Mamm. 220, not of Lesson, 1827, 

which = the typical race from Sumatra. 
1909. Gymnura gymnura minor Lyon, Proc. U.S. Nat. Mus. 36: 453. Trang, 2,000 ft.. 

Lower Siam. 

Genus HYLOMYS MuUer, 1839 

1839. Hylomys Miiller in Temminck, Verb. Nat. Gesch. Nederl. Overz. Bezitt., Zool. 

Zoogd. 50. Hylomys suillus Miiller. 

■, I species : Hylomys suillus, page 1 7 

Hylomys suillus MuUer, 1839 Lesser Gymnura 

Approximate distribution of species: Yunnan (Burmese Border), Burma, Indo- 
China, Siam, Malay States, Tioman Island, Sumatra, Java, Borneo. 



(Hylomys suillus suillcs MuUer, 1839. Extralimital) 

1839. Hylomys suillus Muller in Temminck, Verb. Nat. Gesch. Nederl. Overz. Bezitt., 
Zool. Zoogd. 25, 50. Java. 

Hylomys suillus peguensis Blyth, 1859 

1859. Hylomys peguensis Blyth, J. Asiat. Soc. Bengal, 28: 294. Pegu, Lower Burma. 

Hylomys suillus siamensis Kloss, 191 6 

19 16. Hylomys siamensis Kloss, J.N.H. Soc. Siani, :?.• 10. Hinlap, 900 ft., Eastern 
Siam. Range: to Annam, Laos (Indo-China). 

Hylomys suillus miorotixus Thomas, 1925 

1925. Hylomys suillus microtinus Thomas, P.Z.S. 497. Thai-nien, Tonkin, Indo-China. 

Ranges to Laos, but probably not occurring with the last. Osgood (1932) 

regarded both forms as races of H. suillus. 

Genus NEOTETRACUS Trouessart, 1909 
1909. Neotetracus Trouessart, Ann. Mag. N.H. 4: 389. Neotetracus sinensis Trouessart. 
I species: Neotelracm sinensis, page 18 

Neotetracus sinensis Trouessart, 1909 Shrew-Hedgehog 

Approximate distribution of species: Szechuan and Yunnan, in China; Northern 
Burma; Indo-China. 

Neotetracus sinensis sinensis Trouessart, 1909 

1909. Neotetracus sinensis Trouessart, Ann. Mag. N.H. 4: 390. Tatsienlu, 2,545 m., 
Szechuan, China. Range: Szechuan, Yunnan. 

Neotetr.\cus sinensis fulvescens Osgood, 1932 

1932. Neotetracus sinensis fulvescens Osgood, Field Mus. Publ. Zool. 18: 239. Chapa, 
Tonkin, Indo-China. 

Neotetracus sinensis cuttingi Anthony, 1941 

1941. Neotetracus sinensis cuttingi Anlhony, Field Mus. Publ. Zool. sy: 58. Hpimaw 
Road, above Hpimaw fort, 9,000 ft., North-Eastern Burma. 

Subfamily E r i n a c e i n a e 

Authors arc not yet agreed on how many species of Hedgehogs should be recog- 
nized. For instance, Ognev (1928), in his work on the Mammals of the U.S.S.R., 
recognized four genera containing thirteen species in that region; whereas Bobrinskii 
and Kuzyakin ( 1944) retain in the same region one genus with only four species. 



The late J. L. Chaworth-Musters was for many years doing preparatory work at 
the British Museum for a checklist of Palaearctic Mammals. Through the kindness of 
his executors we have most or aU of his notes in our possession, including the 
synonymy of all the Palaearctic Hedgehogs, and we propose here to adopt most of 
his classification, which retains the three genera which we think it reasonable to 
adopt, and recognizes two or three species in each of them. 

Genus ERINACEUS Linnaeus, 1 758 

1758. Erinaceus Linnaeus, Syst. Nat. loth ed. /.• 52. Erinaceus europaeus Linnaeus. 
1848. Atelerix Pomel, Arch. Sci. Phys. Nat. Geneve, g: 251. Erinaceus albiventris 

Wagner, the Senegambian Hedgehog. Valid as a subgenus. 
1866. Peroechinus Fitzinger, S.B. Akad. Wiss. Wien, ^4: 565, 1866, and ^6: 856, 1867. 

Erinaceus pruneri Wagner from the Sudan. ( =Atelerix.) 
1868. Herinaceus Mina-Palumbo, Ann. Agric. Sicil. 12: 37. {N.V.) (Emendation.) 
1918. Aethechinus Thomas, Ann. Mag. N.H. /.• 194. Erinaceus algirus Duvernoy & 


2 species in the area covered by this list: 
Erinaceus algirus, page 23 
Erinaceus europaeus, page 19 

The first-named belongs to the subgenus Atelerix. It differs from normal species of 
that subgenus in retaining the small hallux which is, however, not constantly sup- 
pressed in the restricted Atelerix of Thomas. In the subgenus Atelerix the prior name is 
E. frontalis Smith, 1831, from South Africa, but E. algirus may be shown to be 
distinct from that by the fact that there is an average size distinction between the two 
species. Thus, often specimens oi E. frontalis examined, only one reaches 52 mm. in 
length of skull; all the others fail to reach 50 mm. But we possess only two specimens 
off. algirus (in a moderate series) which are less than 51 mm. in length. 

Miller, 1912, Cat. Mamm. West Europe, 115, contrasts the characters of the two 
Palaearctic species here admitted. 

Chaworth-Musters used to put all named forms into synonymy, and did not 
recognize any subspecies. Whilst this list is based on his notes, we do not feel that 
such an arrangeinent would be acceptable to the majority of zoologists, and so have 
listed those forms which are likely to be of subspecific value. Bobrinskii divides E. 
europaeus into three groups of races typified by europaeus, roumanicus and amurensis (all 
of which were regarded as species by Ognev). 

Subgenus ERINACEUS Linnaeus, 1758 

Erinaceus europaeus Linnaeus, 1758 European Hedgehog 

Approximate distribution of species: Europe, widely distributed, west to Britain 
and Ireland, north to Norway and Sweden, south to Crete, Greece, Italy, Sicily, 
Spain, and including Denmark, Holland, Belgium, France, Germany, Switzerland, 
Bohemia, Hungary, Yugoslavia, Rumania, Poland, etc. Russia: roughly from north 


palaearc:tic: and Indian mammals i 758-1946 

of Lake Ladoga eastwards, south to the Crimea and Caucasus; Central Siberia in 
part, eastwards roughly to Tomsk district, south to River Emba and North- West 
Kazakstan ; Amur and Ussuri regions in Pacific Siberia. Eastern China : states of 
ChihU, Shensi, Shansi, Hupeh, Shantung, Kiangsu, Anhwei; Manchuria. Asia 
Minor: Palestine. 

Erinaceus europaeus europaeus Linnaeus, 1 758 

1758. Ennacais tiiropaais Linnaeus, Syst. Xat. loth ed. /.• 52. Wamlingbo, South 

Gothland Island, Sweden (see Thomas, 191 1, P.Z.S. 142). 
1779. Hvstrix erinaceus Blumenbach, Handbuch Naturg. 72. Germany. 
1845. Erinaceus caniceps H. Smith, Naturalist's Libr. 1 Jardinc's), 2nd ed. /j; 148. 

Forest of Soignies, near Brussels, Belgium. 
1897. Erinaceus echinus Schulze, Helios Berlin, 14: yi. Substitute for europaeus. 
1900. Erinaceus europaeus occidentalis Barrett-Hamilton, Ann. Mag. N.H. fj: 362. 

Haddington, Scotland. 
1900. Erinaceus europaeus typicus Barrett-Hamilton, loc. cit. 363. 
1912. Erinaceus suillus Miller, Cat. Mamm. Western Europe, 120. France, quoted as 

Geoflroy, Cat. Mammif. Mus. Nat. H.N. 67, but according to a note in 

Chaworth-Musters ALS., this name was nc\'er published by GeofTroy in 

1803; the proofs only are known. 
191 2. Erinaceus caninus Miller, loc. cit. France. Quoted as GeofTroy, Cat. Mammif. 

Mus. Nat. H.N. 68, but according to note in Chaworth-Musters MS. this 

name was never published by GeofTroy in 1803. 
Range: Western Central Europe from Scotland, Southern Norway and Central 
Sweden to Pyrenees and Alps, west to Ireland. 

Erinaceus europaeus conxolor Martin, 1838 

1838. Erinaceus concolor Martin, P.Z.S. iS^y: 103. Near Trebizond, Asia Minor. 

1907. Erinaceus ponticus Satunin, Zool. Anz. 31: 233. Kobuleti, 22 versts north of 

Batum, Georgia, Transcaucasia. 
Range; Transcaucasia, Asia Minor, to Lebanon, Syria (B.M.). 
Ognev regarded this as a species. 

Erinaceus europaeus amurensis Schrcnk, 1859 

1859. Erinaceus europaeus var. amurensis Schrenk, Reisen im Amur-Lande, /, pi. iv. 

fig. 2 : 100-105. Gulssoja, near Aigun, on Amur river. Northern Manchuria, 

(5 Jan. 1859, see verso 2nd title page.) 
1903. Erinaceus orientalis J. .\llen. Bull. Amer. Mus. N.H. ig: 179. Vladivostock, 

Eastern Siberia. 
10(17. Eiiiiineus ussuriensis Satunin, .Ann. Mus. Zool. .Acad. St. Pctcrs'o. 19116, ;/: 

170. Sidemi. Southern Ussuri, Eastern Siberia. 
Range: Korea, Manchuria, South-Eastern Russian Asia. 

Erinaceus europaeus dealbatus Swinhoe, 1870 
1870. Erinaceus dealbatus Swinhoe, P.Z.S. 450. Pekin, C^hihli, China. 
1907. Erinaieus chinensis Satunin, .\nn. Mus. Zonl. Aiail. St. Pctersb. 19116, //: 
173. Tvnl/a-inl/a, Kiiingan .Mnuntains, Manclunia. 


1907. Erinaceus krejenbergi Matschie, Exped. Filchner, Mamm. 135. Type purchased 

in market place, Shanghai, China. 
1907. Erinaceus tschifuensis Matschie, loc. cit. 137. Chefoo, Shantung, China. 

1907. Erinaceus hanensis Matschie, loc. cit. 138. Hankow, Hupeh, China. 

1908. Erinaceus hughi Thomas, Abstr. P.Z.S. 44; 1909, P.Z.S. igo8: 966. Paochi, 

Shensi, China. 
1926. Hemiechinus manchuricus Mori, Annot. Zool. Jap. //.• 108. Koshurei, South 

Manchuria. Status ^^(/p Kuroda. 
Range: China, from Chihh, Hunan, Anhwei, Hupeh, Kiangsu, Shantung, Shensi, 
to Manchuria (part). 

Erinaceus europaeus hispanicus Barrett-Hamilton, 1900 

1900. Erinaceus europaeus hispanicus Barrett-Hamilton, Ann. Mag. N.H. j.- 363. Seville, 
Spain. Range: Iberian Peninsula. 

Erinaceus europaeus italicus Barrett-Hamilton, 1900 

1900. Erinaceus europaeus italicus Barrett-Hamilton, Ann. Mag. N.H. §: 364. Siena, 
Italy. Range: Italy, Ticino in Switzerland, Sardinia. 

Erinaceus europaeus roumanicus Barrett-Hamilton, 1900 

1900. Erinaceus europaeus roumanicus Barrett-Hamilton, Ann. Mag. N.H. 5.- 365. 

Gageni, Prahova, Rumania. 

1901. Erinaceus danubicus Matschie, S.B. Ges. Naturf. Fr. Berlin, 9, 229. Prundu, 

19 15. Erinaceus europaeus roumanicus var. kievensis Charlemagne, Mamm. of the 
neighbourhood of Kiev, 37. {N.V., fide Ognev.) Neighbourhood of Kiev, 
1930. Erinaceus rumanicus rumanicus (morpha) bolkayi \Iartino, Zap. Russk. Nauch. 

Inst. Byelgrad, 2: 60. Cetinje, Montenegro, Yugoslavia. 
1933. Erinaceus roumanicus roumanicus drozdovskii Martino, Prirod. Razpr. 2: 56. 

Kocane, Vardar, Macedonia, Southern Yugoslavia. 
Range: Eastern Germany, Northern Bohemia, Hungary, Rumania, Yugoslavia, 
Greece, Poland, Southern and Central Russia, east to Orenberg and Tomsk Govt. 
in Siberia, south to Crimea, Northern Caucasus. 
Regarded as a species by Ognev and Miller; as a race oi europaeus by Bobrinskii, and 
in synonymy of that species in Chaworth- Musters' MSS. 

Erinaceus europaeus consolei Barrett-Hamilton, 1900 

1900. Erinaceus europaeus consolei Barrett-Hamilton, Ann. Mag. N.H. j.- 366. Near 
Palermo, Sicily. 

Erinaceus europaeus transcaucasicus Satunin, 1905 

1905. Erinaceus europaeus transcaucasicus Satunin, Mitt. Kaukas. Mus. 2: 106, 281. 
' Ordubad on the Araxes river, Transcaucasia. Ognev referred this form to 

roumanicus as a race, and said it occurred in Northern and Southern Caucasus, 

and that the next was probably a synonym. 
1918. Erinaceus roumanicus sacer Thomas, Ann. Mag. N.H. 2: 212. Neighbourhood of 

Jerusalem, Palestine. 



Erinaceus europael's nesiotes Bate, 1906 

1906. Erinaceus europaeus nesiotes Bate, P.Z.S. igo^, ■2: 316. Near Gonia, Western 


Erinaceus europaeus abasgicus Satuniii, 1907 

1907. Erinaceus ponticus abasgicus Satunin, Zool. Anz. ^i: 234. Zebeldinsk part of 

Abchasia on the upper and middle course of River Kodov, \Vestern 
Caucasus. Ognev regarded this as a subspecies oi concolor. 

Erinaceus europaeus .miodon Thomas, 1908 

igo8. Erinaceus miodon Thomas, Abstr. P.Z.S. 44; 1909, P.Z.S. igo8: 965. Yulinfu, 
Shensi, 4,000 ft., China. 

Erinaceus europaeus rhodius Festa, 1914 

1914. Erinaceus europaeus rhodius Festa, Boll. Mus. Zool. Anat. Comp. Torino, zg, 
No. 686, 3. Koskino, Island of Rhodes, Eastern Mediterranean. 

Erinaceus europaeus meridion,\lis Altobello, 1920 

1920. Erinaceus europaeus meridionalis Altobello, Fauna Abruzzo e MoHse, Mamm. i: 
13. Abruzzi, Italy. 

Erinaceus europaeus koreanus Lonnberg, 1922 

1922. Erinaceus koreanus Lonnberg, Ann. Mag. N.H. g: 624. Chosen, Korea. 
(?) 1922. Erinaceus amurensis koreensis Mori, Ann. Mag. N.H. 10: 616. Kaijo, north 
of Seoul, Korea. 

Erinaceus europaeus centralrossicus Ognev, 1926 

1926. Erinaceus europaeus centralrossicus Ogntw, Uchen. Zap. Scv. Kavkaz. Inst. /.• 37. 

Sichevsk, Smolensk Govt., Russia. 
1928. Erinaceus europaeus centralrossicus (natio) pallidus Ognev, Mamm. Eastern 

Europe, Northern Asia, /.• 96. Tyumensk district, Tobolsk Govt., Western 


Erinaceus europaeus dissimilis Stein, 1930 

1930. Erinaceus roumanicus dissimilis Stein, Z. Siiuget, ^: 240. Klcin-Sturlack, Eastern 
Prussia, Germany. 

Incertae scdis 

Erinaceus sibincus Er.xleben, 1777, Syst. Regn. Anini. 172. Siberia (based on Seba, 
1734, Thesaurus, /.• 79, pi. 49, figs. 4, 5). 

Erinaceus or Hemiechinus dauuricus Sundevall, 1842 

This is a very little known species. G. Allen and later Bobrinskii refer it to Hemie- 
chinus; Bobrinskii suggests it may prove to be a well marked subspecies of//, auntus. 
Ognev regarded it as a species oi Erinaceus. A pencil note in Chaworth-Musters' MSS. 
says that A'. dauuricu<, is a genuine Erinaceus, a subspecies of E. europaeus, but that 
Hemiechinus prrj-wahki is //. albulus? G. Allen lists the latter as a synonym oi' dauuricus. 


1842. Erinaceus dauuricus Sundevall, K. Svenska Vetensk. Akad. Handl. 1841: 237. 

Dauuria, Transbaikalia. 
( ?) 1907. He77iiechinus przewalskii Sa.tunm, Ann. Mus. Zool. Acad. St. Petersb. 1906, 

//: 181. Northern China. G. Allen recorded this form from Mongolia, but 

had seen no specimens. 

Subgenus ATELERIX Pomel, 1848 [Synonym: Aethechinus ThomdLi, 1918) 

Erinaceus algirus Duvernoy & Lereboullet, 1842 Algerian Hedgehog 

Approximate distribution of species: Morocco, Algeria, Libya, Canary Islands; 
South-Eastern France, Spain, Balearic Islands. 

Erinaceus algirus algirus Duvernoy & Lereboullet, 1842 

1842. Erinaceus algirus Duvernoy & Lereboullet, Mem. Soc. Hist. Nat. Strasbourg, 

3, 2, art. 1:14. Algeria, no exact locality; Oran given by Miller (191 2) and 

G. Allen (1939). 
1882. Erinaceus fallax Dohson, Monogr. Insectivora, 9. Sfax, Tunisia (type in B.M.). 

Range: Morocco to Libya, Spain, South-Eastern France. 

Erinaceus algirus vagans Thomas, 1901 

1901. Erinaceus algirus vagans Thomas, P.Z.S. igoi, i: 38. San Cristobal, Minorca, 
Balearic Islands. Range includes Majorca. 

Erinaceus algirus caniculus Thomas, 19 15 

1915. Erinaceus algirus caniculus Thomas, Ann. Mag. N.H. 16: 152. Toston, Fuerte- 

ventura Island, Eastern Canary Islands. 
(?) 1877. Erinaceus krugi Peters, S.B. Ges. Naturf. Fr. Berlin, 78. Habitat unknown, 

probably West Africa or Southern Europe. (Type specimen killed in 

Mayaguez, Porto Rico, ? introduced.) 

Erinaceus algirus lavaudeni Cabrera, 1928 

1928. Aethechinus algirus lavaudeni Cabrera, Bol. Soc. Esp. H.N. 28: 454. Mogador, 

Genus HEMIECHINUS Fitzinger, 1866 

1842. Ericius Sundevall, K. .Svenska Vetensk. Akad. Handl. 1841: 223. Erinaceus 

auritus Gmelin. Not of Tilesius, 181 3. 
1866. Hemiechinus Fitzinger, S.B. Akad. VViss. Wien, 5^, i : 565; ibid, 1867, j6: 858. 

Erinaceus platyotis Sundevall = Erinaceus aegyptius Fischer. 
(?) 1928. Erinaceolus Ognev, Mamm. E. Europe, N. Asia, /.• 168. Hemiechinus 

microtis Laptev. 

2 species : Hemiechinus auritus, page 24 
Hemiechinus megalotis, page 26 



Chaworth-Miisters was goina; to retain three species in this genus, H. aurilus, H. 
collaris (to include albulus, turaniciis, minor, alaichankus, persici/s, turfanicus, holdereri, 
major, irisiilaris), and H. rm'galolis. Bobrinskii says all these forms represent one species; 
Ognev further subdivided forms of the genus which occur in the U.S.S.R., and 
retained several more species. The British Museum material gives the following 
cranial measurements: for H. megalotis, length of skull 52-56.9 mm. (average 54 mm., 
two specimens) ; H. collaris as understood by Chaworth-Musters, skull length averages 
about 47 mm. (46.1-48.6 mm.) (based on specimens of collaris from Cutch, Sind, 
Palanpur, Punjab; of albulus from Djarkent, Yarkand, Kashgar, Northern Afghani- 
stan; of turanicus, Transcaspia) ; H. aurilus as restricted by Chaworth-Musters from 
Egy-pt, Palestine, Cyprus, Iraq, Cyrenaica, has the skull length averaging 43.9 mm. 
(42-44.8 mm.). The type of calligoni has also been examined. In our material, there- 
fore, there is an absolute difference in size of skull between the three groups, but in 
Ognev's Key to the Mammals of Eastern Europe, etc., it will be found that there is a 
considerable overlap between auritus and "collaris" as listed by Chaworth-Musters, 
and we think it best to merge these two species, following Bobrinskii. Ognev's form 
major can have the skull as large as in megalotis. Our specimens of megalotis are from 
Kandahar and Baluchistan. 

Hemiechinus auritus Gmelin, 1770 Long-eared Hedgehog 

Approximate distribution of species: Egypt, Cyrenaica; Palestine, Cyprus, Asia 
Minor, Persia, Afghanistan; Punjab, Cutch, Sind, Rajputana, Palanpur in India; 
Chinese Turkestan, Mongolia; Russian Turkestan, where widely distributed north 
to Altai steppe; Caucasus and South-Eastcrn Russia (Volga steppes as far north as 
Kuibyshev, Don steppes). ? Ceylon (B.M., "grayi" is labelled from Wella- 
watte, Ceylon). 

Hemiechinus auritus auritus Gmelin, 1770 

1770. Erinaceus auritus Gmelin, Nov. Comment. Acad. Sci. Petrop. 14: 519. Astra- 
khan, South-Eastern Russia. 

1842. Erinaceus aurilus caspicus Sundevall, K. Svenska Vctensk. Akad. Hand!. 1841, 
237. Emendation of auritus. 

Range: European range of the species and Kazakstan steppes. 

Hemiechinus auritus aegyptius Fischer, 1829 

1829. Erinaceus aegrptius Fischer, Syn. Mamm. 262. Egypt. Based on E. aeg)ptius 

Geoflroyr C;at. .\Ius. H.N. Paris, 1803, which was never published; proof 

sheets only are known ; Chaworth-Mustcrsj. 
1833. Erinaceus libvcus Ehrcnberg in Hemprich & Ehrcnberg, S\'mb. Phys. Mamm. .\- 

sig. k, recto (footnote). Desert near Alexandria, Egypt. 
1842. Erinaceus [Ericius] plalyotis Sundevall, K. -Svenska Vctensk. Akad. Hand!. 1841, 

232. Egypt. 
1882. Erinaceus frontalis Dobson, Monogr. Insect. /.• 18, not of Smith, 1831. See 

Anderson & de Winton, Mamm. Egypt, 1902, 159, as to status. 
Range: Egypt, Cyrenaica. 



Hemiechinus auritus collaris Gray, 1830 

1830. Erinaceus collaris Gray in Hardwicke, Illustr. Indian Zool. /, pi. 8. Doab. 

between the Rivers Jumna and Ganges, India. (See Wroughton, 1910, J, 

Bombay N.H. Soc. 20: 81.) 
1832. Erinaceus spatangus Bennett, P.Z.S. 123. Himalayan mountains. 

1832. Erinaceus gravi 'Qennttt, P.Z.S. 124. Himalayan mountains. 

1833. Erinaceus indicus Royle, Illustr. Bot. Himalaya, 6. Delhi, India. 
Range: Northern India as listed under the species; Afghanistan (B.M.). 

Hemiechinus auritus albulus Stoliczka, 1872 

1872. Erinaceus [Hemiechinus) albulus Stoliczka, J. Asiat. Soc. Bengal, 41, 2: 226. 
Langur, near Sandshu, Yarkand, Chinese Turkestan. 

Hemiechinus auritus syriacus Wood, 1876 

1876. Erinaceus syriacus ^Vood, Bible Animals, 83. Palestine. This name is available if 
the Palestine form is recognizable. 

Hemiechinus auritus calligoni Satunin, 1901 

1 90 1. Erinaceus calligoni Satunin, Prot. Obshch. Est. Kazan, No. 192 (misprinted 

191), 2. P.Z.S. igoi, 2: 284. Village of Aralyk, about 40 versts south of 

Erivan, Armenia. Range: Daghestan, Transcaucasia. 

Hemiechinus auritus turanicus Satunin, 1905 

1905. Erinaceus albulus turanicus Satunin, Mitt. Kaukas. Mus. 2: 45, 70. Ferghana, 

Usbekistan, Russian Turkestan (see Satunin, 1906, Ann. Mus. Zool. Acad. 
- St. Petersb. //.• 180.) Range: from Kopet-Dag to Lake Balkash and 

Semirechyia (Ognev). 

Hemiechinus auritus minor Satunin, 1907 

1907. Hemiechinus albulus minor Satunin, Ann. Mus. Zool. Acad. St. Petersb. 1906, 
//: 180. Barnaul, W'estern Siberia. 

Hemiechinus auritus alaschanicus Satunin, 1907 

1907. Hemiechinus albulus alaschanicus Satunin, Ann. Mus. Zool. Acad. St. Petersb. 
1906, //: 181. Alashan, Inner Mongolia. 

Hemiechinus auritus persicus Satunin, 1907 

1907. Hemiechinus persicus Salunin, Ann. Mus. Zool. Acad. St. Petersb. 1906, //: 184. 

Guljander, Persia. 

Hemiechin'us auritus brachyotis Satunin, 1908 

1908. Hemiechinus calligoni brachyotis Satunin, Mitt. Kaukas. Mus. 4: 47, 106. Ach-su, 

Semacha district, Transcaucasia. 

Hemiechinus auritus turfanicus Matschie, 191 1 

191 1. Hemiechinus albulus turfanicus Matschie in Futterer, Durch Asien, 5, 5, Zool.: 4 
(of reprint). Chami, Sinkiang (Chinese Turkestan). 


Hemiechinus auritus holdereri Matschie, 1922 

1922. Hemiechinus holdereri Matschie, S.B. Ges. Naturf. Fr. Berlin, 73. Near River 
Sarin-gol, Gobi Desert, Mongolia. 

Hemiechinus auritus major OcjneN- cS: Heptncr, 1028 

1928. Hemiechinus alhulus major Ognev & Hcptner, Zool. Anz. 75; 259. Station 
Annau, Ashabad, Transcaspia, Russian Turkestan. 

Hemiechinus auritus turkestanicus Ognev, 1928 

1928. Hemiechinus calligorii turkestanicus Ognev, Mamm. E. Europe, N. Asia, /.• 130. 
Station Kara-Usyak, north of Perovsk, Russian Turkestan. 

Hemiechinus auritus insularis Timofejcw, 1934 

1934. Hemiechinus alhulus insularis Timofejcw, Zool. J. Moscow, ij: 748, 758. Island 
of Barsa Kelmes in the Sea of Aral, Russian Gentral Asia. 

Hemiechinus megalotis BIyth, 1845 Afghan Hedgehog 

Approximate distribution of species; Baluchistan, Afghanistan and South-\\'estern 
Russian Turkestan. 

Hemiechinus megalotis BIyth, 1845 

1845. Erinaceus megalotis BIyth, J. Asiat. Soc. Bengal, i^: 353 ifootnote). Kandahar, 

(?") 1926. Hemiechinus chorassanicus Laptev, Bull. Uniw Asie Cent, ij: 1 15-1 16. \'alley 

Tchandyr, near Atrek river, Kopet-Dagh, South-Western Russian Turkestan. 

Bobrinskii suggests this is a cross between H. auritus and Paraechinus hvpo- 

?nela5; Chaworth-Musters regarded it as a synonym of//, megalotis. 

Incertae sedis 

Hemiechinus russowi Sntimn, 1907, Ann. Miis. Zool. .\cad. St. Petersb. 1906, //: 177. 
Tchinaz, Samarkand district, south of Tashkent, Russian Turkestan. The 
skull is unknown. According to Bobrinskii it "docs not even represent a very 
pronounced individual aberration". 

Hemiechinus microtis Laptev, 1925, Bull. Univ. Asie Cent. 8: 66. Tashkent, Russian 
Turkestan. Type of Ennaceolus Ognev. "Only known by two specimens 
from Tashkent, we regard as a pronounced aberration, all the distinctive 
features of which are connected with an anomalous under-development 
of the organ of hearing" (Bobrinskii & Kuzyakin). 

Hemiechinus homalacanthus Stroganov, 1944, C;.R. Acad. Sci. U.R.S.S. 44. ■',: 120. 
Kabadiani, Tadjikistan, Russian Turkestan. From descriptions a large form, 
perhaps representing //. megalotis. 

Genus PARAECHINUS Trouessart, 1879 

1870. Paraechinus Trouessart, Key. Zool. Paris, j: 242. Erinaceus micropus BIyth. 
1907. Macroechinus Satunin, Ann. Mus. Zool. Acad. St. Petersb. 1906, //: 189. 
Eiinaceii^ krpnmelas 13randt. 



3 species: Paraechinus aethiopicus, page 27 
Paraechiniu hypomelas, page 28 
Paraechinus micropus, page 28 

There are two distinct groups in this genus, typified by hypomelas and aethiopicus. 
Chaworth-Musters was going to retain three species {micropus was the third) but did 
not deal with the Madras form, nudiventris . We have two skulls for the last-named 
which have the zygoma incomplete and apparently lack the jugal, but it does not 
seem a constant character as the zygoma is complete in a third specimen. Chaworth- 
Musters' species may be retained and keyed as follows: nudiventris is tentatively 
referred to micropus, and micropus might well be regarded as an eastern representative 
of aethiopicus. 

1. P 3 less reduced, three-rooted. Skull appears long and narrow, narrow in ptery- 

goid region (pterygoid width averages about 51 per cent, or less of length of 
skull). Paraechinus hypomelas 
P 3 single-rooted (occasionally two-rooted), but very reduced. Skull appears 
wider, and is wider in pterygoid region (pterygoid width averages 54 per cent, 
or more of length of skull). 2 

2. Pterygoid width very rarely under 25 mm. (only once in fourteen specimens). 

Paraechinus aethiopicus 
Pterygoid width normally less than 25 mm. (fifteen out of sixteen specimens). 

Paraechinus micropus 

On South-\Vestern Asiatic and North African forms (in part) see Morrison-Scott, 
1939, Novit. Zool. 41: 202. 

Paraechinus aethiopicus Ehrenberg, 1833 Ethiopian Hedgehog 

Appro.ximate distribution of species: Morocco, Algeria, Asben, Sudan, Arabia, 

(Par.'\echinus aethiopicus aethiopicus Ehrenberg, "1833. Extralimital) 

1833. Erinaceus aethiopicus Ehrenberg in Hemprich. & Ehrenberg, Symb. Phys. 

Mamm. 2: sig. k, recto (footnote). Dongola Desert, Sudan. 
1839. Erinaceus sennaariensis Hedenborg, Isis, 32.- 8, nom. nud. 
1841. Erinaceus brachydactylus Wagner, Schreber Saugeth. Suppl. 2: 24. Renaming of 

1867. Hemiechinus pallidus Fitzinger, S.B. Akad. Wiss. ^Vien, ^6, 1: 866. Senaar, 


Paraechinus aethiopicus deserti Loche, 1858 

1858. Erinaceus deserti Loche, Cat. Mamm. Oiseaux Algerie, 20. Southern Sahara, in 
Oasis of Beni-Mzab, Ouargla, and Tuggurt, Algeria. Ranges to Morocco. 

Paraechinus aethiopicus pectoralis Heuglin, i85i 

1861. Hemiechinus pectoralis Heuglin, Nov. Acta. Leop. Carol. 2g: 22. Petra, Trans- 

c 27 


Paraechinus aethiopicus dorsalis Anderson & de VVinton, 1901 

1901. Erinaceus dorsalis Anderson & de VVinton, Ann. Mag. N.H. y: 42. Hadramaut, 
Southern Arabia. 

Par.'^echini's aethiopicus LUDLOW! Thomas, 191 9 

1919. Paraechinus hidlowi Thomas, J. Bombay N.H. Soc. 26: 748. Hitt, on the 
Euphrates, about 100 miles west of Baghdad, Iraq. 


1 92 1. Paraechinus deserti blancalis Thomas, Ann. Mag. N.H. 8: 570. Island of Djerba, 

South-Eastern Tunis. 

Paraechinus .aethiopicus albatus Thomas, 1922 

1922. Paraechinus dorsalis albatus Thomas, Ann. Mag. N.H. g: 144. Tanb Island, 

Persian Gulf. 

Par.\eciiinus aethiopicus oniscus Thomas, 1922 

1922. Paraechinus oniscus Thomas, Ann. Mag. N.H. 10: 307. Fayush, 7 miles north of 
Sheikh Othman, near Aden, Southern Arabia. 

Par.-vechinus aethiopicus albior Pocock, 1934 

1934. Paraechinus dorsalis alhior Pocock, Ann. Mag. N.H. 14: 636. Dhimir Wad, 
Geradun, 960 ft., Southern Arabia. 

Paraechinus micropus Blyth, 1846 Indian Hedgehog 

.Approximate distribution of species: Punjab, Rajputana, Sind, Cutch, Kathiawar, 
Palanpur and Madras, India. 

P.-vraechinus micropus micropus Blyth, 1846 

1846. Erinaceus micropus Blyth, J. Asiat. Soc. Bengal, i§: 170. Bhawalpur, Punjab, 

Northern India. 
1867. I/emiechinus inentalis Fitzinger, S.B. Akad. W'iss. W'ien, 56", i : 874. (Gray, 1843, 

Cat. Mamm. B.M. 81, nom. nud.). ? Himalayas. 
(?) 1872. Erinaceus [Hemiechinus) pictus Stoliczka, J. Asiat. Soc. Bengal, 41, 2: 223. 

VVestern part of Cutch, India. 
Range: as in the species, except Madras. 

Paraechinus (?) micropus .\udiventris Horsficld, 1851 

1851. Erinaceus nudivenlris Horsfield, Cat. Mamm. .\Ius. E. India Co. 136. Madras, 
Southern India. 

Paraechinus hypomelas Brandt, 1836 Brandt's Hedgehog 

Approximate distril ■' irm of species: Russian Turkestan (Ust-L'rt, Turkmcnia, 
Usbekistan as far north as Samarkand Province); Persia, Afghanistan, Arabia; Sind 
Punjab (Salt Rantjo and North- West Frontier (Peshawar). 


Paraechinus hypomelas hypomelas Brandt, 1836 

1836. Erinaceus hypomelas Brandt, Bull. Sci. St. Petersb. /; 32. Northern Persia. 

(See Ognev, 1927, Zool. Anz. 6g: 210-212.) 
1875. Erinaceus macracanthus Blanford, Ann. Mag. N.H. 16: 310. Near Kerman 

(Carmania), 5,000-6,000 ft., Persia. 
1918. Paraechinus amir Thomas, Ann. Mag. N.H. /.• 232. Kandahar, Afghanistan. 
Range: Persia, Afghanistan, Russian Turkestan. 

Paraechinus hypomelas blanfordi Anderson, 1878 

1878. Erinaceus blanfordi Anderson, J. Asiat. See. Bengal, ^7, 2: 208. Rohri, Sind, 

North-Western India. 
1878. Erinaceus jerdoni Anderson, loc. cil. 209. Karachi, Sind, India. 

Paraechinus hypomelas nicer Blanford, 1878 

1878. Erinaceus wz^cr Blanford, J. Asiat. Soc. Bengal, ^7, 2: 212. Muscat, Arabia. 

Paraechinus hypomelas seniculus Thomas, 1922 

1922. Paraechinus niger seniculus Thomas, Ann. Mag. N.H. g: 142. Island of Tanb, 
Persian Gulf. 

Paraechinus hypomelas sabaeus Thomas, 1922 

1922. Paraechinus niger sabaeus Thomas, Ann. Mag. N.H. g: 143. El Kubar, about 
60 miles north of Aden, 5,200 ft., Arabia. 

Paraechinus hypomelas eversm.^'iNni Ognev, 1927 

1927. Paraechinus hypomelas eversmanni Ognev, Zool. Anz. 6g: 218. Ust-Urt, east of 
Caspian Sea, Northern Russian Turkestan. 


Genera: Desmana, page 32 
Galemys, page 33 
Scapanulus, page 35 
Scaptonyx, page 34 
Talpa, page 35 
Uropsilus, page 31 
Urotrichus, page 33 

On these genera see Cabrera (1925). Another helpful work is W'inge, 1923, 
Pattedyr Slaegter, i: 143-155 (key, 154-155). About a dozen genera are currently 
recognized in this family in the Old World which Simpson, following Thomas and 
Cabrera, divides into four subfamilies. It is beginning to be understood that no useful 
purpose is served by -recognizing genera based solely on dental formulae in this 
family. Thus Schwarz (1948) refers all members of the subfamily Talpinae to one 
genus, in one species of which are four different dental formulae which have hitherto 



been considered as of generic \alue. Similarly, Osgood ! 1937) has shown conclusively 
that in the UropsiUnae the three supposed genera of Thomas are of very little value, 
being based solely on the presence or absence of vanishing teeth which give three 
supposedly different dental formulae.' For this family we are fortunate in possessing 
the manuscript which Chaworth-Musters prepared for a list of Palaearctic Mammals. 
The subfamilies and genera here admitted may be separated as follows: 

1. The upper canine is the dominant front tooth; it is conspicuously larger than the 

incisors in front of it. Animal modified for underground life. Tail very short. 
Hand very large, larger than in other Asiatic and European genera, the inner 
side conspicuously broadened. (.Subfamily T.\lpin.\e) TALPA 
The upper canine is not the dominant front tooth, but the first upper incisor is 
generally very strongly so. In one genus (.Scaplonvx) none of the front teeth are 
much enlarged. 2 

2. .\nimal modified for aquatic life; hindfeet very broad and large; tail long, nearly 

as long as, or longer than, head and body, at least partly specialized for swim- 
ming. First upper incisor very large. (Subfamily Desmani.xae) 3 

Animal not aquatic; tail not specialized for swimming, and hindfeet less 
broadened. 4 

3. Tail flattened laterally throughout; unicuspid teeth low and thick; ridges on 

braincase unusually developed; head and body 180-215 mm. {Ognev), tail 
1 70-2 1 5 mm. DESMANA 

Tail flattened laterally only at end; unicuspid teeth slender; ridges on braincase 
moderate; head and body circa i 10-156 mm., tail circa 126-156 mm. 


4. Animal shrew-like; tail long, sometimes as long as head and body, and usually 

over 80 per cent, of it, poorly haired; hands small, not fossorial. First upper 
incisor dominant but not very strongly enlarged. Head and body length under 
90 mm. (Subfamily Uropsilinae) UROPSILUS 
Animal mole-like; tail well haired, often almost bushv, short, averaging 55 per 
cent, at most of head and body, but more often less than 40 per cent, of it. 
Hands large and broad, fossorial; but less broadened, particularly on inner side, 
than in Talpinae. (Subfamily Scvlopinwe) 5 

^ Precisely similar conditions occur in the .African family Chrysochloridac. .'Vs many as nine genera 
have been admitted, and no two authors agree which are valid and which are of subgeneric value 
when endeavouring to make revision: nor will they do so until they realize that presence or absence 
of vanishing teeth are of not much value. Thus in ^^Neambhsomus^'' three quite different formulae are 
found in the same series; and Chrysochloris as restricted by Roberts, for which a large series has been 
collected in the West Cape, has the formula varying indi\'idually so that three formulae at least can 
be present. One of us (J. R. E.) has examined the great majority of type specimens in this family, and 
inclines to the view that there are only three main generic types in this family: Chrysospalax. the giant 
golden-moles with the posterior zygoma root and occiput much enlarged ; Chrysochloris^ containing the 
majority of the subgenera and species, small animals with posterior zygoma root and occiput normal, 
and two functional fmgers in the hand; and Eremiialpa, like Chrysochloris^ but hand with three 
functional fmgers. Surely in animals so highly modified for digging as these arc, the latter character 
is \'ery much more important than any dental formula.'* 



First upper incisor not much enlarged; 42 teeth, 11 upper, 10 lower. (Head and 

body length 90 mm., and less.) SCAPTONIX 
First upper incisor very enlarged; 36 or 38 teeth. 6 

Head and body length roughly 100 mm. (98-108 mm.). Nine upper and 9 lower 

Head and body length very rarely reaches 100 mm. (four only in eighty-five 
specimens noted in B.M.). Ten upper, 8 or 9 lower teeth. UROTRICHUS 

Subfamily Uropsilinae 

Genus UROPSILUS Milne-Edwards, 1872 

1872. Uropsilus Milne-Edwards in David, Nouv. Arch. Mus. H.N. Paris, 7; Bull. 92. 
Uropsilus soricipes Milne-Edwards. 

191 1. Nasillus Thomas, Abstr. P.Z.S. 49; P.Z.S. 129. Nasillus gracilis Thomas. 

1912. Rhynchonax Thomdii, P.Z.S. 1^0. Rkynckonax andersoni Thomas. 

I species : Uropsilus soricipes, page 3 1 

Three genera, based on three dental formulae which are now known not to be 
constant, are tentatively referred to a single species. (It may be noted that in Talpa 
micrura as recently defined by Schwarz, four different dental formulae occur.) See 
particularly Osgood, 1937, Field Mus. Publ. ^ool. 20, 27: 365. G. Allen thought that 
the three groups should stand as genera until they can be shown to be not generically 
valid, and argues that the three groups are probably distinct as they have fairly 
distinct areas of geographical distribution. Against this it might be argued that as 
they do not occur together they are probably all races of one species. The three are 
hardly distinguishable from each other externally. Osgood retained two genera, but 
his diagnosis is not very convincing, and he apparently thought Rhynchonax andersoni 
was a race of U. soricipes, while the other two named forms of Rhynchonax, atronates 
and nivatus, he thought might be races of Nasillus gracilis. Until the contrary is 
proved we prefer to retain one species only, which is considered as on the point 
of losing some small teeth, so that different individuals may either have them 
or not. 

Uropsilus soricipes Milne-Edwards, 1872 Shrew-Mole 

Appro.ximate distribution of species: Szechuan and Yunnan in China, to Northern 

Uropsilus soricipes soricipes Milne-Edwards, 1872 

1872. Uropsilus soricipes Milne-Edwards, Nouv. Arch. Mus. H.N. Paris, 7, Bull.: 92. 
Rech. H.N. Mamm. 1872, 272. Moupin, Szechuan, China. 


Uropsilus soricipes gracilis Thomas, 191 1 

191 1. Aasilliis gracilis Thomas, Abstr. P.Z.S. No. 100, 49. igi2, P.Z.S. 130. Mt. 

Chinfusan, near Nanchwan, 4,000 ft., Szechuan, China. Osgood (1937) 

retains this form as a species. 

Uropsilus soricipes .\ndersoni Thomas, 191 1 

1911. Rhynchonax andersoni Thomas, Abstr. P.Z.S. No. 100, 49. /g/i', P.Z.S. 130. 

Omisan, Omei Hsien, Southern Szechuan, 9,500 ft., China. 
1923. Rhmchonax andersoni atronates G. Allen, Amer. Mus. Novit. No. 100, 2. Mucheng 

Sahveen drainage, South-Western Yunnan, 7,000 ft., China. Status fide 

Anthony, 1941, Field Mus. Publ. Zool. 27: G2. But O.sgood thought it might 

be a race oi gracilis. 
Range: to Northern Burma. 

Uropsilus soricipes investig.^tor Thomas, 1922 

1922. Nastllus investigator Thomas, Ann. Mag. N.H. 10: 393. Kiukiang-Salween 

divide at 28" N., Yunnan, 1 1,000 ft., China. 

Uropsilus soricipes nivatus G. Allen, 1923 

1923. Rhynchonax andersoni nivatus G. Allen, Amer. Mus. Novit. No. 100, 2. Ssu-shan 

(Snow Mountain), Likiang Range, Western Yunnan, 12,000 ft., China. 
(Osgood thought this might be a race oi gracilis.) 

Subfamily Desmaninae 

Genus DESMANA Guldenstacdt, 1777 

1777. Desrnana Guldenstacdt, Beschaft Berl. Ges. Naturf. Fr. 3: 108. Castor moschatus 

1799. Desman Laccpede, Tabl. Mamm. 7. Castor moschatus Linnaeus. 

1800. Mrgale Cuvier, Leq. Anat. Comp. i, Tabl. i. Castor moschatus Linnaeus. 
1815. Desmanus Rafincsque, Analyse de la Nature, 59. Renaming oi Mygale. 

1829. Mxogalea Fischer, Synops. Mamm. 250. Substitute for Mygale. 

1830. C.apiios ^Vagler, Nat. Syst. Amphib. 14. Substitute for Alygale. 
1836. .\/)Offa/c Brandt, Wiegmann's Arch. Naturgcsch. /.• 176. 

I species: Desrnana moschata, page 32 

Desrnana moschata Linnaeus, 1758 Russian Desman 

.Ali[jn,ximatc distribution of species: "The basins of the \'olga, Don and Mius. 
East to the lower Kama, north to upper Unzha; west to Ruibinsk, the confluence of 
the .Mi'sha with the X'olga, Moscow and Kharkos-, and south to the Don (it does not 
(«( ur (in the 1< ft-ljank tributaries of that river) and X'olga deha, and the middle and 
lM\\<r Ri\cr Ural" Tjubrinskii). 



Desmana moschata Linnaeus, 1758 

1758. Castor moschatus Linnaeus, Syst. Nat. loth ed. /; 59. Russia. 
181 1. Mygale moscovitica Geoffroy, Ann. Mus. H.N. Paris, ly: 192. Substitute for 
moschatus Linnaeus. 

Genus GALEMYS Kaup, 1829 

1829. Galemys Kaup, Skizz. Europ. Thierwelt, /.• 119. Mygale pyrenaica Geoffroy. 
1835. Mygalina I. Geoffroy in Gervais, Resume des Legons de Mamm. 45. Adygale 

pyrenaica Geoffroy. 
1846. Galomys Agassiz, Nom. Zool. Index Univ. 159. Emendation oi Galemys. 

I species: Galemys pyrenaicus, page 33 

Galemys pyrenaicus Geoffroy, 181 1 Pyrenean Desman 

Approximate distribution of species: Pyrenean region of South- Western France, 
Spain and Portugal (see Puissegur, 1937, Recherches sur le Desman des Pyrenees, 
Bull. Soc. H.N. Toulouse, 6j: 163-225, map, 2 pis., 104 figs.). 

Galemys pyrenaicus pyrenaicus Geoffroy, 181 1 

181 1. Mygale pyrenaica Geoffroy, Ann. Mus. H.N. Paris, ly: 193. Near Tarbes, 
Hautes-Pyrenees, France. Range: to North-Eastern Spain. 

Galemys pyrenaicus rufulus Graells, 1897 

1897. Myogalea rufula Graells, Mem. R. Accad. Madrid, ly: 460. Rio Balsain, above 
the Venta de los Mosquitos, Sierra de Guadarrama, Segovia, Central Spain. 

Subfamily Scalopinae 

Three genera in Asia, aU apparently closely allied to each other, are admitted in 
this typically Nearctic subfamily. 

Genus UROTRICHUS Temminck, 1841 

1839. Urotrichus Temminck, Tijdschr. Natuur. Gesch. j: 286. Urotrichus talpoides 

Temminck, nom. nud. 
1841. Urotrichus Temminck, Het. Instit. K. Ned. Inst. 212. Urotrichus talpoides 

1887. Dymecodon True, Proc. U.S. Nat. Mus. 1886: 97. Dymecodon pilirostris True. 

2 species : Urotrichus pilirostris, page 34 
Urotrichus talpoides, page 34 

U. pilirostris was named as a distinct genus Dymecodon, characterized by having nine 
lower teeth (two lower incisors), thereby differing from typical Urotrichus which has 
eight lower teeth (one lower incisor). Bearing in mind that within Uropsilus and Talpa 
as here understood and as defined in part by Schwarz and by Osgood, different 



dental formulae occur in the same species, and also bearing in mind Simpson's state- 
ment that animals from similar localities are likely to be allied to each other, this is 
not a character of even subgeneric value. However, the tail seems about half head 
and body length in pilirostris so far as can be at present ascertained, whereas in 
talpoides it is normally below 40 per cent, of that measurement, and this character 
combined with the extra lower tooth suggests that here it is possible that we are 
dealing with two valid species. 

Urotrichus talpoides Temminck, 1841 Japanese Shrew-Mole 

Approximate distribution of species: Japan (apparently Hondo southwards). 

Urotrichus talpoides talpoides Temminck, 1841 

1 84 1. Urotrichus talpoides Temminck, Het. Instit. K. Ned. Inst. 215. Nagasaki, 

Kiushiu, Japan. (See Kuroda, 1938, List. Jap. Mamm. 87.) 
1906. Urotrichus talpoides pilirostris Thomas, P.Z.S. /5105, 2: 342. Not of True, 1886. 

Urotrichus talpoides adversus Thomas, 1908 

1908. Urotrichus talpoides adversus Thomas, P.Z.S. 49. Sasuna, North Island, Tsushima 
Islands, Japan. 

Urotrichus t,\lpoides centralis Thomas, 1908 

1908. Urotrichus talpoides centralis Thomas, P.Z.S. 50. Jinrio, Tokushima Ken, 500 ft., 
Shikoku Island, Japan. 

Urotrichus talpoides hondonis Thomas, 1908 

1908. Urotrichus talpoides hondonis Thomas, P.Z.S. 51. Nakaomi, near Ohitu, Izo, 

Hondo, Japan. 
1929. Urotrichus talpoides yokohamanis Kanda, Zool. Mag. Tokyo, ^i: 147. [N.V.) 

Yokohama, Hondo, Japan. 

Urotrichus talpoides minutus Tokuda, 1932 

1932. Urotrichus talpoides minutus Tokuda, Annot. Zool. Jap. /j.- 580. Dogo Island, 
Oki Islands, Japan. 

Urotrichus pilirostris True, 1886 True's Shrew-Mole 

Approximate distribution of species: Hondo, Japan. 

Urotrichus pilirostris True, 1886 

1886. Dymecodon pilirostris True, Proc. U.S. Nat. Mus. g: c^y. Enoshima (Venosima), 
at mouth of Bay of Veddo, Hondo, Japan. 

Genus SCAPTONYX .\Iilnc-Edwards, 1872 

1872. Scaptonyx Milne-Edwards in David, Nouv. Arch. Mus. H.N. Paris, y: Bull. q-2. 
Scaptonvx fusicauda David. 

I species: Scaptonvx Jusicaudus, page 35 


Scaptonyx fusicaudus Milne-Edwards, 1872 Long-tailed Mole 

Approximate distribution of species: Szechuan and Yunnan in China; Northern 

Scaptonyx fusicaudus fusicaudus Milne-Edwards, 1872 

1872. Scaptonyx fusicauda Milne-Edwards in David, Nouv. Arch. Mus. H.N. Paris, y: 

Bull. 92. Borders of Kukunor and Szechuan, China. 
1872. Scaptonyx fusicaudatus Milne-Edwards, Rech. H.N. Mamm. 278. Borders of 

Kukunor and Szechuan, China. 

Scaptonyx fusicaudus affinis Thomas, 19 12 

1912. Scaptonyx fusicaudatus affinis Thomas, Ann. Mag. N.H. g: 514. Twelve miles 

south-east of Atunsi, North-Western Yunnan, 13,500 ft., China. Range: 

Yunnan, Northern Burma. 

Genus SCAPANULUS Thomas, 191 2 
19 1 2. Scapanulus Thomas, Ann. Mag. N.H. 10: 396. Scapanulus oweni Thomas. 
I species: Scapanulus oweni, page 35 

Scapanulus oweni Thomas, 19 12 Kansu Mole 

Approximate distribution of species: China, States of Kansu, Szechuan and 

For notes on this genus see also G. Allen, 1938, Mamm. China & Mongolia, i: 81. 
The Nearctic Neiirotrichus has a similar dental formula. W't have few specimens for 
either, but our Scapanulus has a much larger hand, and thicker, hairier tail than our 
Neiirotrichus, and the first upper incisor seems larger in Scapanulus. 

Scapanulus oweni Thomas, 191 2 

1912. Scapanulus oweni Thomas, Ann. Mag. N.H. 10: 397. Twenty-three miles south- 
east of Taochou, Kansu, 9,000 ft., China. 

Subfamily T a 1 p i n a e 

For revision, see Schwarz, 1948, Revision of the Old World Moles of the genus 
Talpa, P.Z.S. 118: 36-48. 

Genus TALPA Linnaeus, 1 758 

1758. Talpa Linnaeus, Syst. Nat. loth ed. /.■ 52. Talpa europaea Linnaeus. 

1848. Mogera Pomel, Arch. Sci. Phys. Nat. Geneve, g: 246. Talpa wogura Temminck. 

1867. Scaptochirus Milne-Edwards, Ann. Sci. Nat. Zool. y: 375. Scaptochirus moschalus 

1875. Parascaplor Gill, Bull. U.S. Geo). & Geogr. Surv. Terr. /, 2: no. Talpa leucura 

1898. Chiroscaptor Heude, Mem. H.N. Emp. Chin. 4, i : 36. Chiroscaptor sinensis 

Heude = Scaptochirus moschatus Milne-Edwards. 
1940. Euroscaptor Miller, J. Mamm. 21: 443. Talpa klossi Thomas. 



Talpa 'contd.] 

1941. Eoscalops StroganoN-, C.R. Acad. Sci. URSS. 33: 270. Talpa longirostris Milne- 
1941. Asiosialops Strosanov, Cl.R. Acad. Sci. URSS. 33: 271. Talpa allaka Nikolsky. 
1948. Asioscaplor Schwarz, P.Z.S. 118: 36. Error for Asioicalops Stroganov. 

3 species: Talpa caeca, page 38 

Talpa europaea, page 37 

Talpa mkrura, page 39 
It is \cry difficult to decide how many species should be retained in this genus. 
Miller ( 191 2) recognized four in Europe, europaea, caeca, romana and occidenlalis, and in 
the latest revision of the genus (Schwarz, 1948) this classification is followed exactly. 
Ognev in his work on the Mammals of the U.S.S.R. also retained four species, 
europaea, caeca, allaka and caucaska, but his characters were not very convincing, and 
Schwarz, while retaining altaka, makes caucaska a synonym oi europaea. More recently 
Bobrinskii and Kuzyakin refer all forms from the U.S.S.R. to a single species europaea, 
with groups of races typified by europaea, caeca and altaica. Chaworth-Musters' manu- 
script agrees with Bobrinskii's arrangement for the Western and Central Asiatic 
members of the genus. Against this it must be stated that the range of T. europaea 
overlaps that of T. caeca in Switzerland and apparently in the Caucasus. On account 
of this, the latter is here listed as a valid species. There is an average size difference 
between the two, caeca being the smaller. Schwarz, who seems to oversplit the western 
section of the genus, does the reverse with the Eastern Asiatic forms. AU of these he 
refers to a single species for which the prior name is Talpa mkrura. Hitherto these 
have been distributed among four genera, Talpa, Mogera, Parascaptor and Scaptochirus, 
aU of them based on dental formulae which Schwarz shows are not constant. The 
nosepad is long, naked and grooved on the upper side in T. mkrura as understood by 
Schwarz, the penis is said to be speciaUzed (on this account .Miller separated those 
forms which retain the primitive 44 teeth as Euroscaptor), and the tail is much 
shortened; in B.M. material this is usually 20 mm. or less in length, except for the 
large form kobeae in which it averages only 14 per cent, of the head and body. The 
western species have the tail very rarely as short as 20 mm. T. allaica approaches the 
micrura group, apparently, in the structure of the nosepad, and the tail is relatively 
short (although in Bobrinskii's figures it is rarely under 20 mm.). This author states 
that the europaea moles in the U.S.S.R. are small, but with large teeth and a long tail, 
while the altaka moles are considerably larger, but with small teeth and a short tail, 
but that the two groups are connected by intermediate forms and, excepting in the 
Caucasus, all conform to a definite law; as one goes south, and particularly east, the 
size of the animals increases while their teeth and tail become smaller. He gives 
figures to support this. So that although Schwarz says that altaka is definitely not 
europaea, we suggest that as this form does not occur together with europaea it might, 
f .llowing Bobrinskii and Chaworth-Musters, be considered a very distinct repre- 
sentative race. It seems not \cry much more distinct from europaea than some forms, 
notably moschatus, which Schwarz refers as a subspecies to T. micrura, are from the 
latter. If in the future subgeneric division is required for the micrura group, then 
Mogera is thr prior name. 


Talpa europaea group. 

Talpa europaea Linnaeus, 1 758 Common Mole 

Approximate distribution of species: Europe, widely distributed; north to Southern 
Sweden; south to Spain, Italy, Sicily and Northern Greece; west to Britain; east to 
Ural Mountains and Caucasus. Occurs in France, Belgium, Holland, Denmark, 
Finland, Poland, Germany, Switzerland, Transylvania, Yugoslavia, Rumania, 
Bulgaria. According to Kuzyakin and Bobrinskii, represented in the Siberian Altai 
region of Lake Baikal, Lena river, near Yakutsk, Northern Yenesei — apparently 
extending north of the Arctic Circle and to Mongolia. 

Talpa europaea europaea Linnaeus, 1 758 

1758. Talpa europaea Linnaeus, Syst. Nat. loth ed. /.• 52. Engelholm, Kristianstad, 
Southern Sweden. (Chaworth-Musters' MSS. See Skanska Reise, 352, mis- 
printed "332" in Linnaeus, loc. cit. 52.) 

1772. Talpa caudata Boddaert, Kortbegrip Nat. /.• 50. (jV.F.) Renaming oi europaea. 

1776. Talpa frisius Muller, in Linnaeus, Natursyst. Nat. Suppl. 36. East Friesland, 


1777. Talpa europaea albo-maculata Erxleben, Syst. Regn. Anim. /.■ 117. East Fries- 

1785. Talpa vulgaris Boddaert, Eiench. Anim. /.• 126. Renaming of europaea. 
1788. Talpa europaea alba Gmelin, Linn. Syst. Nat. 13th ed. /.• no. Sweden. 
1788. Talpa europaea cinerea Gmelin, loc. cit. Eifel, Germany. 
1788. Talpa europaea variegata Gmelin, loc. cit. Sweden. 
1792. Talpa europaea nigra Kerr, Anim. Kingd. 200. Renaming oi europaea. 
1797. Talpa europaea ruja Borkhausen, Der Zoologe fCompendiose Bibliothek 

gemeinn. Kenntn. f. alle Stande, 21) Heft. 5-8: 13. (A'.L., teste Miller). 

Southern France. 
(?) 1800. Talpa europaea major Bechstein in Pennant, Allgem. Uebers Vierf. Thiere, 

2: 725. Siberia, no exact locality. 
1836. Talpa europaea flavescens Reichenbach, Der Naturfreund, figs. 472-3. Saxony, 

1852. Talpa europaea albida Reichenbach, VoUstand. Naturgesch. .}: 336. Germany. 
1852. Talpa europaea lutea Reichenbach, loc. cit. Germany. 
i86g. Talpa europaea maculata Fitzinger, S.B. Akad. W'iss. Wien. 551, i : 401. Renaming 

of albo-maculata. 
1869. Talpa europaea grisea Fitzinger, loc. cit. 403. .Synonym of cinerea wrongly attri- 
buted to Zimmermann, 1780. 
1897. Talpa scalops Schulze, Helios Berlin, /^.' gi. Renaming of europaea. 
1908. Talpa europaea brauneri Satunin, Mitt. Kaukas. Mus. ^: 2, 8. Post Cuculi, 

Belitsk district, Bessarabia. 
1908. Talpa coeca caucasica Satunin, Mitt. Kaukas. Mus. ^.' 5-q. Stavropol, Caucasus. 

(Status_^rff Schwarz.) 
1925. Talpa europaea uralensis Ognev, Bull. Soc. Nat. Moscou, jjjj, 1-2: 4. District of 

Perm, Russia. 
1930. Talpa europaea pancici Martino, Zap. Russk. Nauch. Inst. Byelgrad, 2: 60. 

Kraljevo, Serbia, Yugoslavia. 



Talpa europaea europaea [conld.] 

1 93 1. Talpa romana stankovici Martino, J. Mamm. 12: 53. Magerevo Mountains, 

Perister, Macedonia, 1,000 m., Southern Serbia. 
Range: European range of the species, except Sicily; in Italy, south to Tuscany; in 
Russia, north to the region of the White Sea (absent from Crimea). 

T.\LPA (?) EUROPAEA ALTAICA Nikolsky, 1 883 

1883. Talpa altaica Nikolsky, Trans. Soc. Nat. St. Pctersb. 14: 165. Valley of the 

Tourak, Altai Mountains, Siberia. 
1905. Talpa coeca var. suschkini Kastschenko, Trans. Tomsk. Univ. 27: 75 (of reprint). 

Sayan Mountains, Central Siberia. 
1921. Talpa altaica saianensis Bielovusev, Ann. Mus. Zool. Acad. St. Petersb. 22: 

xviii. Kazir-Susko Forest, Sayan Mountains, 2,000 ft., Siberia. 
(?) 1922. Talpa europaea var. irkiilensis Dybowski, Arch. Nauk. Biol. Lwow, /, 6-8: 4 
[rioni. niid.). Irkutsk, Siberia. 

1936. Talpa altaica salauica Egorin, Trav. Inst. Sci. Biol. Tomsk, 2: 154. Salair 

Mountains, Tomsk Govt., Siberia. 

1937. Talpa altaica tymensis Egorin, Trav. Inst. Sci. Biol. Tomsk. 4: 49. Tymsk, 

Naunak, on River Vasyugan, tributary of River Ob, Siberia. 
1937. Talpa altaica sibirica Egorin, Trav. Inst. Sci. Biol. Tomsk, 4: 51. Avseenko, 

Tyazhin, near Mariinsk, Western Siberia 
Range: Asiatic range of the species, above. 
Schwarz considers this a distinct species. 

Talpa europaea romana Thomas, 1902 

1902. Talpa romana Thomas, Ann. Mag. N.H. 10: 516. Ostia, near Rome, Italy. 

1920. Talpa romana major Altobello, Fauna Abruzzo e Molise, Mamm. /.• 32. 
Abruzzi, Italy. Not of Bechstein, 1800. 

1925. Talpa romana monlana Cabrera, Genera Mamm. 87. Mm. nov. for major Alto- 
bello, preoccupied. 

Range: Italy and Sicily. 

Schwarz considers this a distinct species, chiefly characterized by large teeth and 
dental details. It is not known to occur with europaea, and Chaworth-Musters 
treated it as europaea. 

Talpa europaea ognevi Stroganov, 1944 

1944. Talpa romana ognevi Stroganov, C.R. Acad. Sci. U.R.S.S. 44, "i: 121. Bakuriana, 

Georgia, Transcaucasia. 
The status of the next is not sure. Neither this nor apparently the last were 
allocated by Schwarz. 

1945. Talpa europaea traiiscaiicasica Dahl, Zool. Pap. Biol. Inst. Erevan, 3, 48. {M.V.). 

\'(iskrescnkovva, Kirovakan, Armenia. 

Talpa caeca S.ivi, 1822 Mediterranean Mole 

.\ppro.\imatc distribution of species: Portugal, Spain, Switzerland, Italy, Yugo- 
sKi\ia, Greece, ,'\sia Minor, Caucasus. Treated as a subspecies of europaea by 
Bobrinskii and Kuzyakin, and in synonyiny ui' europaea by Clhaworth-Musters, but it 
occurs with europaea in several places in Switzerland, and in Caucasia. Averages 
smaller in size europaea. 



Talpa caeca caeca Savi, 1822 

1822. Talpa caeca Savi, Nuovo Giorn. de Letterati, Pisa, /; 265. Near Pisa, Italy. 
1884. Scaptochirus davidianus Milne-Edwards, C.R. Acad. Sci. Paris, gg: 1 143. Said to 
have come from borders of Syria and Asia Minor. Not of Swinhoe, 1870. 

1906. Talpa coeca levantis Thomas, Ann. Mag. N.H. ij: 416. Scalita, south of 

Trebizond, Asia Minor. 

1925. Talpa hercegovinensis Bolkay, Nov. Mus. Sarajevo, No. 1:1. Stolac, Herzegovina, 


1926. Talpa coeca orientalis Ognev, Uchen Zap. Sev. Kavkaz. Inst. /.• 33, 55. Chosta, 

Black Sea Govt., Southern Russia. 
1932. Talpa olympica Chaworth-Musters, Ann. Mag. N.H. g: 166. Eastern slope 

Mount Olympus, Thessaly, 800 m., Greece. 
Range : Switzerland and Italy to Asia Minor and Caucasus. 

Talpa caeca occidentalis Cabrera, 1907 

1907. Talpa caeca occidentalis Cabrera, Ann. Mag. N.H. 20: 212. La Granja, Segovia, 

Spain. Range: Spain and Portugal. Schwarz gives this form specific rank. 

Talpa micrura group. 
The classification of Schwarz, 1948, is followed. 

Talpa micrura Hodgson, 1841 Eastern Mole 

Approximate distribution of species, as understood by Schwarz: from Ussuri 
region of South-Eastern Siberia, Manchuria, Korea, Japan, Formosa, Eastern 
MongoHa {fide Schwarz), the greater part or all of the major states of China (Kansu 
apparently excepted) ; to Indo-China, Siam, Malay States, and Burma, Assam, west- 
wards to Sikkim and Nepal. 

Talpa micrura micrura Hodgson, 1841 

1 84 1. Talpa micrurus Hodgson, Calcutta J. N.H. 2: 221. Nepal, Central and Northern 

1843. Talpa cryplura Blyth, J. Asiat. Soc. Bengal, 12: 177. Sylhet, Assam. 
1858. Talpa macrura Hodgson, J. Asiat. Soc. Bengal, 2j: 176. Near Darjeeling, 

7,000 ft., India. (Status _/?rft! Schwarz.) 
Range: Nepal, Sikkim, Assam. 

Talpa micrura wogura Temminck, 1842 

1842. Talpa wogura Temminck, in Siebold's Fauna Japonica, Mamm. /.• ig. 

Nagasaki, Kiushiu, Japan. 
1845. Talpa moogura Temminck, loc. cit. 4: tab. 4, figs. 1-5. Misspelling oi wogura. 
1880. Talpa mizura Gunther, P.Z.S. 441. Neighbourhood of Yokohama, Japan. 
1936. Mogera wogura minor Kuroda, Botany & Zoology, Tokyo, 4, i: 74. Shiobora, 

Pref Tochigi, Central Hondo, Japan. 
1936. Mogera wogura gracilis Kishida, Nikko No. Shokubutsu to Dobutsu, 261. 

{N.V., ? nom. nud.). Near Shobugahama, Nikko, Japan. 
Range: Japan, including Hondo, Shikoku, Oki Islands. 



Talpa micrura leucura Blyth, 1850 

1850. Talpa leucura Blyth, J. Asiat. Soc. Bengal, ig: 215, pi. 4, fia;. i. Cheriapuuji, in 
Khasi Hills, Assam. 

(?) 1929. Talpa klossi Thomas, Ann. Mag. N.H. j: 206. Hue Nya Pla, 10 miles north- 
west of Raheng, 2,500 ft., Siam. 

(?) 1940. Talpa parvidens Miller, J. Mamm. 21: 203. In forest at agricultural station 
of Blao, near the upper Donai River, Annani, Indo-China. (See Schwarz, 
1948: 46.) 

Range: Assam, Burma, Siam, Laos, Clochin-Clhina, Annam, ? Yunnan, and Malay 
(Some lines have accidentally been omitted from Schwarz's paper in dealing with 

this race.) 

Talp.x micrura insul.'\ris Swinhoe, 1862 

1862. Talpa insularii Swinhoe, P.Z.S. 356. Formosa. 

Talpa micrura moschata Milne-Edwards, 18G7 

1867. Scaplochints moicfiatiis Milne-Edwards, Ann. Sci. Nat. Zool. y: 375. Swanhwafu, 

100 miles north-west of Pekin, Chihli, Clhina. 
1870. Scaptochirus davidiarms Swinhoe, P.Z.S. 620. Accidental renaming oi moschatus. 
1881. Talpa h'ptura Thomas, Ann. Mag. N.H. y: 470. Neighbourhood of Pekin, 

Chihli, China. 
1898. Chiroscaptor sinensis Heude, Mem. H.X. Emp. Chin. 4: 36. South-Eastern 

C;hihli, China. 
1898. Scaptochirus moschiferus Heude, lac. cit. 40. Accidental renaming of moschatus. 
1910. Scaptochirus gilliesi Thomas, Ann. Mag. N.H. 5.- 350. Ho-tsin, South-Western 

.Shansi, China. 
1941. Parascaplor grandidens Strogano\', C.R. Acad. Sci. U.R.S.S. jj: 271. Tuntzia- 

Intza (Tunchia Yingtze), east of Dolon Nor, Southern Khingan Mountains, 

Jehol, North-Eastern China. 
Range: C;hihli, Jehol, Shansi, Shensi, Shantung, in China. 

T.\lpa micrura longirostris Milne-Edwards, 1870 

1870. Talpa longirostris Milne-Edwards, C.R. Acad. Sci. Paris, jo: 341. Moupin, 

Szechuan, China. 
1907. Mogera latouchei Thomas, P.Z.S. 463. Kuatun, Fukien, 3,500 ft., South-Eastern 

1940. Euroscaptor grandii Miller, J. Mamm. 21: 444. Mt. Omei, 5,000 ft., Szechuan, 


Talpa mk:rura rorusta Nehring, 1891 

1 89 1. Mngera robusta Nehring, S.B. Ges. Naturf. Fr. Berlin, No. 6: 95. \'ladi\'ostock, 
Eastern Siberia. Range: Ussuri region, Manchuria. 

Talp.\ mi(::rura koheae Thomas, 1905 

1905. Mogera woguia kobcac Thomas, Ann. Mag. N.H. /j.' 487. Kobe, Hondo, Japan. 



Talpa micrura kanai Thomas, 1906 

1906. Alogera wogura kanai Thomas, P.Z.S. /505, 2: 361. Miyanoura, Yakushima, 

1938. Mogera wogura kiusiuana "Kishida", Kuroda, List Jap. Mamm. Tokyo, 89, 

nom. nud. 
Range includes Tsushima, Kiushiu and the Goto group, Japan. 

Talpa micrura coreana Thomas, 1907 

1907. Mogera wogura coreana Thomas, P.Z.S. 463. Kim-hoa, 65 miles north-east of 

Seoul, 300 ft., Korea. 

Talpa micrura hainana Thomas, 1910 

1910. Mogera hainana Thomas, Ann. Mag. N.H. §: 535. Mt. Wuchi, Island of 
Hainan, China. (Apparently not dealt with by Schwarz.) 


Genera: Anourosorex, page 87 
Blarinella, page 55 
Chimmarogale, page 87 
Crocidura, page 70 
Diplomesodon, page 86 
Feroculus, page 86 
Nectogale, page 88 
Neomys, page 61 
Solisorex, page 86 
Sorex, page 43 
Soriculus, page 56 
Suncus, page 64 

According to Simpson (1945), so far as the present region is concerned, the above 
genera fall into two subfamilies: the Soricinae, with Sorex, Blarinella, Soriculus and 
Neomys; and the Crocidurinae with the remainder. 

This division into subfamilies, which is adopted by many authors, seems to be 
based chiefly on the Soricinae having the teeth pigmented at the tips, and the 
Crocidurinae having the teeth entirely white. Other authors, e.g. Allen, Miller and 
Bobrinskii, do not recognize these subfamilies and we concur with them, especially 
in view of the variability in the extent, or even the entire absence, of the pigmentation 
of the teeth in the Soricinae. 

The Soricidae is perhaps the most difficult of all mammalian families to deal with, 
so far as the species are concerned. Chaworth-Musters was attempting the task just 
before his untimely death, but the only manuscript he left deals with some of the 
Palaearctic species of the genus Sorex. 



The twelve genera represented in London may be distinguished as follows : 

1. Teeth with the cusps pigmented, red or brown; pigmentation can become very 

weak but is normally traceable. (Subfamily Soricinae of Simpson)' 2 

Teeth all white. (Subfamily Crocidurinae of Simpson) 5 

2. Slightly modified for aquatic life; the hindfeet large and fringed; the tail long, 

its underside keeled or hairy. First lower incisor more or less simple, scarcely 
lobed. ' NEOMYS 
No aquatic modification. 3 

3. First lower incisor simpler, with only one prominent lobe on its cutting edge. 

First lower incisor more complex, with more than- one lobe on its cutting edge, 
usually three or four traceable. 4 

4. The last two upper unicuspid teeth excessively minute, hardlv traceable. 

The last two upper unicuspids quite well marked. SOREX 

5. Externally modified for underground life; tail scarcely apparent externally, 

shorter than hindfoot. Ears much reduced. Seven upper teeth; M 3 vestigial. 

Externally not much modified for burrowing; tail clearly longer than hindfoot. 
Except Diplomesodon, more than 7 upper teeth. G 

6. Considerably modified for aquatic life; tail long, hairy, more or less fringed 

below. Ear small or absent. 7 

Not modified for aquatic life. 8 

7. Much more specialized for aquatic life; toes fully vvebbed; no external ear trace- 

able; tail with fringes of hair each side and above and below. Braincase very 
wide. NECTOGALE ' 

Much less specialized for aquatic life; toes not webbed; tail hairy, only the 
underside slightly fringed; with external ear. CHEMMAROGALE 

8. Colour piebald, quite distinct from all other Soricidae examined 'below, sides, 

cheeks and a patch in the middle of the back, white; otherwise the back grey, 
but much white showing on the sides) ; tail hairy, tufted, and white. Soles 
slightly hairy. Seven upper teeth; M 3 not vestigial. DIPLOMESODON 

Colour not as just described. More than 7 upper teeth. 9 

g. Foreclaws strongly enlarged. 10 

Foreclaws not enlarged. 11 

10. First lower incisor more complex, with several lobes on its cutting edge (as in 
Sorex) ; upper teeth (30 teeth in all) ; clear elongated bristles on the tail (such 
as are characteristic 1 if most of the species (A Crocidura and Suncw.). 

First lower incisor simple; 8 upper teeth : 28 teeth in all); no elongated bristles 
on the tail. SOLISOREX 

1 Pii^iiipntation nf tcrth often weak in Soncului and almost untraceable in the two named Formosan 
forms of that genus. 



II. 30 teeth (4 upper unicuspids). SUNCUS 

28 teeth (3 upper unicuspids). CROCIDURA 

Genus SOREX Linnaeus, 1758 

1758. Sorex Linnaeus, Syst. Nat. loth ed. /.• 53. Sorex araneus Linnaeus. 

1829. Oxyrhin Kaup, Entw. Gesch. u. Nat. Syst. Europ. Thierwelt, /.• 120. Sorex 

tetragonurus Hermann (see Miller, 191 2, 29). 
1835. Amphisorex Duvernoy^ Mem. Soc. Mus. H.N. Strasbourg, 2: 23. [Sorex 

hermanni Duvernoy = Neomys fodiens skull, plus Sorex araneus tetragonurus 

1838. Corsira Gray, P.Z.S. 123. Sorex vulgaris = Sorex araneus Linnaeus. 
1842. Otisorex De Kay, Zool. of New York, /; Mamm. 22. Sorex platyrhinus = Sorex 

personatus Geoffrey, from North America. 
1890. Homalurus Schulze, Schriften Nat. Vereins Harzes in \Vernigerode, 5.- 28. 

Sorex alpinus Schinz. 
1927. Soricidus A\Xoht\\o, Rev. Franc. ALimm. /.■ 6. Soricidus monsvairani AXtoheWo = 

Sorex araneus tetragonurus Hermann. (See Gulino, 1939, Boll. Mus. Zool. 

Anat. Comp. Torino, 4y: 136.) 

Apparently 9 species in the Palaearctic region: 
Sorex alpinus, page 54 
Sorex araneus, page 50 
Sorex buchariensis, page 54 
Sorex caecutiens, page 48 
Sorex cylindricauda, page 55 
Sorex daphaenodon, page 53 
Sorex hawkeri, page 46 
Sorex minutus, page 47 
Sorex pacificus, page 54 

This genus is exceedingly difficult to classify, and at 'the present day there are 
nearly a hundred named forms in the Palaearctic. The greatest number of species 
occur in the U.S.S.R. Ognev, 1928, Mamm. U.S.S.R., recognized nineteen in that 
country, but more recently Bobrinskii and Kuzyakin (1944) give a more compressed 
classification of Russian Sorex in which only half a dozen species are retained. These 
authors consider that the large number of named forms is due to a lack of knowledge 
of individual, seasonal and age variations, and their classification incorporates a 
tentative assessment of these. Chaworth-Musters did not complete his manu- 
script for the classification of Sorex, but the first fact that emerges from it is that the 
species now widely known as Sorex macropygmaeus Miller, 1901, must be called 
Sorex caecutiens Laxmann, 1788 [Nova Acta Acad. Set. Pelrop. j: 285). Among 
forms which Bobrinskii and Kuzyakin would reduce to subspecific rank or less, 
Chaworth-Musters was going to retain as species S. daphaenodon, S. raddei, S. shinto 
and S. tundrensis (the latter typically from North America, with various Asiatic 

D 43 


Tlic classificatimi (if Kuzvakin and I'obriiiskii lor the U.S.S.R. was as follows: 

1. (:(lnd^•lobasal Icii^th of skull imt rxcccciini; i)..' niiii. I.cnyth of upper tooth- 

idw under ;,.!! mm. Widtli oi skull nut UKjre than 7 mm. Tail short, not more 
than 31 mm. Hindliiot (without claws) generally not o\'er 8.7 mm. Sorex 
tic/icrskii OEjne\', 19 13, here considered referable to Sorex hmvkcri Thomas, ir)o6. 
C'ond\liibasal length of skull nut less ih.m 14. M mm. Length (il Upper toothrow 
over () nuu. Width of skull o\er 7 lum. Length of tail nut less than 33 mm. 
HindlDot (i\er 9 mm. 2 

2. Width of skull not more th.ui ?■, mm. Total length of skull in large majority of 

cases under i(Li miu., cundylobasal length not more than 17.3 mm. (usually 
under iIlj mm.). Sccrmd upper intermediate tooth markedly smaller than the 
third or 1 less frequenth) the same size. Hindloot usualh less than 11 mm. 

Sorex mi?uitiis 
Width of skull not less than 8.2 mm. Total length of skull over 16.3 mm.Condylo- 
basal length not less than 1(3.5 mm. Second upper intermediate tooth markedly 
larger than third or con\ersely smaller than it. Hiiidfoot ox'cr 10.5 miu. 3 

3. Second upper intermediate tooth considerably smaller than third. Total length of 

skull 17.J mm. Length of upper toothrow 7.!) mm. Sorex biicharicnsis 
Known from one specimen f lund in the North- Western Pamirs. In size and 
configurati<in of skull it is like .S'. macropvgmacus ( = caeculicns) but in structure 
of the teeth it differs from all shrews in the Palaearctic.) 
.Second upper intermediate tooth luarkedly larger than third. 4 

4. Head and body not more than 84 lum. Hindfoot less than 16 imn. Condylobasal 

not more than 21 mm. L'pper toothrow length under 9.5 mm. Distance be- 
t\vecn antrorbital f iraiuina not luore than 3.5 lum. Fourth intermediate tooth 
of upper jaw the same size as, or markedly smaller than, third intermediate 

tooth. 5 

Head and body 87 mm. Hindfoot 17 mm. C)ondylobasal length of skull 23.3 mm. 
Upper toothrow 10 nun. Distance between anteorbital foramina 4.3 mm. 
fourth u|3prr intermediate tooth larger than third. Snrcx jmcijiciis 

(Described from Oregon, U.S.A. To this species Bobrinskii and Kuzyakin 
refer the Ussuri f uni mirabilis: "Comparing .S'. mirahilis with the excellent 
photographs of skulls, measurements and descriptions oi S. pacijiciis gi\en in 
H.Jackson's monograph of the American shrews, 1928, we have been unable 
to find a siirgle feature by whi( h the L^ssuri shrews ma\- \\ith certainty be 
distinguished from the form S. p. Jiaiificiis") 

■-,. C!ond\lol3asal length ol skull in large majorit)' of cases under 18 mm. Length of 
upper toothrow not more than 8 mm. Anteorbital foramina close together, 
distanie lietween them ncjt more than 2.8 mm. 

Siirt'x macrofn'smaeus Miller, 190 1 = Sorex cairiilirns Laxmaiui, 1788 
flondvlobasal length of skull over 18 inm. Length of upper toothrow in large 
majority of cases not less than 8 miu. (usually considerably more). Distance 
between anteorbital foramina in most cases over 2.8 mm. Sorex araneus 


According to notes in Bobrinskii and Kuzyakin, there is a wide individual variation 
to be found in forms which have been regarded as distinct species but which they 
refer to S. araneus. They state, for instance, that the hindfoot length in the Caucasian 
form [satunini] is 1 1.6- 12 mm., whereas in S. a. pnicinius the hindfoot without claws 
reaches 15 mm. The body length even in one form [tomensis) varies between 53 and 
84 mm. The colour pattern varies from almost complete uniformity (in such forms as 
isodon, raddei, unguiculatus, nithenus) through the common two-coloured type to the 
markedly three-coloured type in which the light colouring of the flanks reaches the 
back. The three forms are connected by a great number of transitional stages, and 
not isolated geographically. Again, flat-skulled individuals (such as platjcranius, 
thomasi, turuckanensis, iochanseni) are distributed in the same places as specimens with a 
normal braincase. The relative sizes of the small intermediate teeth are also said to 
vary individually in this species. Bobrinskii and Kuzyakin state that in S. caecutiens 
koreni alone three different types of colouring have been noted, and that coloration is 
useless for diagnostic purposes. These authors incline to ignore all named Russian and 
Siberian subspecies in the two widely ranging allied species, S. araneus and S. caecutiens. 

Chaworth-Musters told us that in his opinion Bobrinskii had "lumped" too far in 
the Sorex of the U.S.S.R., particularly as regards S. daphaenodon, which was described 
as having an unusually hairy tail and heavily pigmented teeth, and which he con- 
sidered a very distinct species, and in deference to his opinion that species is here 
retained, although externally it is not separable from .S'. araneus as here understood. 

Miller (1912) recognized three species of the genus in \V'cstern Europe, as follows: 

1. Anterior lower incisor with low, sometimes ill-defined lobes on cutting edge; first 

lower unicuspid two-pointed ; lachrymal foramen over point of contact between 
M I and IVI 2. Tail about as long as head and body. Sorex alpinus 
(This species is confined to Central Europe, and does not occur in Russia.) 
Anterior lower incisor with high, distinct lobes on cutting edge; first lower uni- 
cuspid single-pointed; lachrymal foramen in front of point of contact between 
M I and M 2. Tail shorter than head and body. 2 

2. First, second and third unicuspids subequal; condylobasal length of skull 14.8- 

16.6 mm. Head and body about 50-60 mm. Sorex minulus 

First and second upper unicuspid much larger than third; condylobasal length of 
skull 17.4-20 mm. fiead and body usually about 65-80 mm. Sorex araneus 

It appears to us from Miller's cranial measurements that the great majority of 
specimens of the last-named have the condylobasal length vcit seldom under 18 mm. 
(cf Bobrinskii's characters for the species), except the Spanish race granarius, which 
surely represents .S'. caecutiens? Few, if any, of the species outside Europe, except the 
striped S. cylindricauda have the tail as long as S. alpinus in B.M. material. 

G. Allen, 1938, Alamm. China & Mongolia, retained half a dozen species from 
this region, as follows: 

I. Back uniform shade of brown without black median stripe. 2 

Back with blackish median stripe. Sorex cylindricauda 

(\Vhich has from Allen's measurements the greatest length of skull, 16.6- 
18.5 mm., and is a tropical species.) 


pai..\i:arc;tic: and indlw mammals 1758-1916 

■2. Lara;cr, hindfiiot with claws 13-14 mm. 3 

Smallor, hindl'oot with claws 12 mm. or less. 5 

3. Lower surfaces whitish-tipped. 4 

Lower surfaces distincth- brownish. Sorex sinalis 

4. Tail about 40 mm. Sorex arancits 
Tail about 50 mm. Sorex excelsus 

5. Skull length about 18 mm. Sorex buxloni Allen, 1903 =- .S'. macropjgmaeus Miller, 

iC)Oi,Jide Kuzyakin and Bobrinskii = Sorex eacciilien^ L.ixmann, 1788. 
.Skull lencjth about 15 mm. Sorex miniitiis 

Chaworth-Musters was Roing to list excehus and i/««//j as distinct species; but if one 
follows the arrangement of Bobrinskii and Kuzyakin, certainly iinalis and probably 
excelsus might be regarded as outlying forms of .S'. araiieus. 

In Lidia this genus almost fails to occur. Only S. cMindricaucla comes into Northern 
Burma, and Miller described a form (planieeps) from Kashmir which in all probability 
represents .S'. minulin. In South- Western Asia there are one or two very early (perhaps 
imidcntifiable) names from Persia; Bodenheimer listed both .S'. araneus and .S'. minutus 
from Palestine; and Thomas named a form from Asia Minor which Bobrinskii and 
Kuzyakin refer to S. araneus. 

The listing of this genus must of necessity be regarded as provisional. 

Sorex ha\vkeri Thomas, 1906 

Approximate distribution of species: Russia, part; Siberia, from approximately 
region of Lake Baikal and the Nizhnaya Tunguska River, eastwards to Kamtchatka, 
Sakhalin, Ussuri and Nijni Kolymsk; Northern Mongolia (Bobrinskii ; Japan. 

Sorex hawkeri H.-wyKERi Thomas, 1906 

1906. ,S'o;r.v //rtirAc/; Thomas, P.Z.S. /5/05, 2: 339. Inukawa, Ycdo, Hondo, Japan. 

Sorex hawkeri Tsr:HERSKii Ognev, 191 3 

1913. Sorex heherskii Ognev, Ann. Mus. Zool. Acad. St. Pctersb. 18: 412. Odarka, 

Lake C:h.iuka, Ussuri region, Eastern Siberia. 
Kuzyakin and Bobrinskii state that owing to lack of material it is not possible to 
give descriptions (jf the geographical variation. The folowing are named from the 
U.S.S.R. which <irc referable to this species. 
191 5. .S'o;r.v /'«)/;( 17 Thomas, Ann. Mag. N.H. /j.' 499. Listvineechnoya, near Irkutsk, 

Lake Baikal, 1,400 ft., Siberia. 
ir)2i. Siiiex neglectus Ognev, Ann. Mus. Zool. Acad. St. Petersb. r:?.- 324. 

Tesovo forest, Mozhaysk district, Russia. 
1921. Sorex itMurieirus Ognev, loc. cit. 326. Valley of Ri\er Bikin, Ussuri region, 

Last( in Siberia. "Given adequate material, it would be good tn determine 

whrilirr or not the features of \Sorcx usuiriemi.i' fall outside the limits of 

inrli\ \ ari.ition" (Bobrinskii and Kuzyakin). 
1933. .Son.v in\iiiieii\n ezekanovskii NaumofT, Abstr. Zool Inst. Moscow' Univ. /.■ 72. 

Till a, Lnwer Tunguska Ri\cr, Clentral Siberia. 



Sorex minutus Linnaeus, 1766 Pysniy Shrew 

Approximate distribution of species: Britain and Ireland, Norway and Sweden, 
France, Germany, Holland, Denmark, Hungary, to Transylvania, Switzerland, 
Italy, Greece, Poland; forest and forest-steppe zones of Russia, Siberia and the Far 
East as far as the Shartar Islands and Sakhalin, and including the Caucasus; 
Szechuan and Tsaidam; North Kurile Islands; apparently Kashmir; Palestine {fide 

Miller, 191 2, Cat. Mamm. W. Europe, recognized two races in Western Europe: 


1766. Sorex minutus Linnaeus, Syst. Nat. 12th ed. /; 73. Barnaul, Western Siberia 

(based on Laxmann's MS. oi Sibir. Briefe). 
1769. Sorex pygmaeus Laxmann, Sibir. Briefe, 72. Barnaul, Siberia. 
1780. Sorex minutissimus Zimmermann, Geogr. Gesch. 2: 385. Yenesei River, Siberia. 
1788. Sorex exilis Gmelin, in Linn. Syst. Nat. 13th ed. /." 1 15. Yenesei River, Siberia. 
1806. Sorex canaliculatus Ljungh, K. Svenska Vetensk. Akad. Handl. 2y: 263. Lom- 

maryd Vigorage, Northern Vedbo district, Jornkoping, Sweden. 
181 1. Sorex minimus GeofTroy, Ann. Mus. H.N. Paris, ly: 186. Selo Kiiskoe, between 

Tomsk and Atchinsk, Siberia. 
1832. Sorex pumilio Wagler, Isis, 2^: 54. Bavaria, Germany. 
1838. Sorex rusticus }enyns, Ann. N.H. /." 423. Near Cambridge, England. 
1838. Sorex rusticus var S[orex) hibernicus Jenyns, loc. cit. Dublin, Ireland. 
1844. Sorex pumilus Nilsson, K. Svenska Vetensk. Akad. Handl. /.• 33. North-Eastern 

Skaane, Sweden. 
1928. Sorex minutus minutus natio melanderi Ognev, Mamm. E. Europe & N. Asia, 

/.• 245. Smolensk Govt., Russia. 
Range: Siberia, Russia, European range of species except Southern Italy and Greece. 

Sorex minutus lucanius Miller, 1909 

igog. Sorex minutus lucanius Miller, Ann. Mag. N.H. 5.- 417. Monte Sirino, Lagonegro, 

Since Miller, the following forms have been named from AVestern Europe: 

1932. Sorex minutus gyvinurus Chaworth-Musters, Ann. Mag. N.H. g: 167. Eastern 

slope Mt. Olympus, Thessaly, 800 m., Greece. 
1940. Sorex minutus insulaebellae Heim de Balsac, C.R. Acad. Sci. Paris, 211, 11: 213. 

Belle Isle, Western France. 

Kuzyakin & Bobrinskii, 1944, Mamm. U.S.S.R., seem to regard the next two 
named forms as valid. 

Sorex minutus gmelini Pallas, 181 1 

181 1. Sorex gmelini Pallas, Zoogr. Ross. As. /.• 134, pi. 10, fig. 3. Crimea (Ognev). 
(This name is used by both Bobrinskii and Ognev, but Chaworth-Musters 
in his synonymy of the species stated: "ig28. Sorex minutus gmelini Ognev, 
Mamm. E. Europe, N. Asia, /.■ 251. Crimea, not Sorex gmelini Pallas, 



1907. Sonw mimttus <;racilliiiiiis Thomas, P.Z.S. 408. D.iriiu', j-, miles norlh-wcst of 
Kiirsaknir, Sakli.iliii Island. Also rccnidcd Hokkaido and Kmca. 

Other named forms Ik mi the U.S.S.R., apparenth nut rec;aided as valid by 
Kuzy.ikin and Bobrinskii. are: 

1 02 1. Soiiw minulus volnuchini Ognev, Ann. Mus. Zool. Acad. St. Petersb. ri\- 32'2. 

Kisha River, Kouban region, \orth- Western Caucasus. 
192'v Sorcx minutiis (morpha) kastchenkoi }ohi\n?,en. Trans. Tomsk Uni\-. js: 66. 

Novii-kusk, C^hulim River, Tomsk district, Siberia. 

C'.hincsc and Indian forms apparently representing this species: 

SoREX MixuTLS THiBF.rAMis Kastschcnko, 1905 

1905. Sorex minutiii thib(tanus Kastschenko, Trans. Univ. Tomsk, 2y: 93 (of reprint). 
Tsaidam, Chinese Central Asia. Also recorded from Szcchuan. China. 

Sorex (.') minutus planiceps Miller, 191 1 

191 1. Sorex planiceps Miller, Proc. Biol. Soc. Washington, i;^: 242. Dachin, Kishtvvar, 
().ooo ft.. Kashmir. 

Sorex (?) minutus leucogaster Kuroda, 1933 

1933. Sorex IcHcogasler ¥^\\roA^, Bull. Biogcogr. Soc. Jap. j, 3: 155. Nasanki, Amamu- 

shiru, 200 ft., North Kurile Islands. 
(?) if|30. Sorex vamaihiiiai Kishida, Z. Mag. Tiik\-o, XLII, 373, nom. run/. 

.Sorex (?J minutus hyojironis Kuroda, 1939 

it)39. Sorcx araneus hvojiroiiis Kuroda, Bull. Biogeogr. Soc. Tokyo, 9; 40. J.damute, 

east of Hailar, Northern Manchuria. The measurements in the description 

suggest a very short-tailed form oi' minutus. 

Sorex caecutiens T..i\mann, 1788 Laxmann's Shrew 

Ap|3r(ixiniatc distributicm of species: apparently Spain and Sweden; Irom Baltic 
Republics and Karelia, Finland, east through the U.S.S.R. to the Chukotski 
Peninsula, Kamtchatka and Sakhalin, north to the Arctic coasts; the southern limit 
of the range runs from C^cntral Ukraine to Gorki Province, thence by the upper River 
Uial .ind the .Altai thrnugh M(jng(ili.i In Kurea, Kansu .md fapcUi. 

.SoRix oAijjrriExs cAKcrriENs Laxmann, 1788 

1700. Surex caeculieiis Laxmann, Xii\. .Vcta Acad. Sci. Petrcip. ijSj, 3: 28-,. l')\ Lake 

Baikal ? = ncighbourhoud nl Irkutsk, Siberia. Hitherto knriwn as S. niiicro- 

/'Vl^maeui, fnit according tii Chawdrth-Mustcrs' MS. this much earlier name 

is valid. 

In the U.S.S.R., Bobiinskii .md Ku/y.ikin do nut recngnizc subspecies ol ,S'. 

m(ieiii/ii:;in(ieu\ -- caicutiens, but the\" refer .V. huxloni, S. iiniiexm, S. haikaletisn. S. tinuniin, 

S. JiDilii .md .v. iillni/in. .ill "I whiili li.i\i- been considered distini t species, to the 

[Hisent spi-( ii-s. 



Chaworth-Musters in his MS. reii.ined shinto as a distinct species, and referred the 
forms ultunus, petschorae and middendorffi as races to Sorex tundrensis Merriam, 1900, Proc. 
Washington Acad. Sci. 2: 16, St. Michael's, Alaska. 

Russian and Siberian forms in order of naming are : 

1901. Sorex macropygmaeus Miller, Proc. Biol. Soc. Washington, i^: 158. Pctro- 
paulski, Kamtchatka. (Synonym 1933. Sorex macropygmaeus macropygmaeus 
natio tungussensis Naumofl", Abstr. Zool. Inst. Moscow Univ. /.• 72. Lower 
Tunguska River, Turukhansk region, North-Western Siberia.) 

1903. Sorex buxtoni ]. Allen, Bull. Amer. Mus. N.H. ig: 181. Gichiga, west coast 
Okhotsk Sea, Siberia. (Synonym, according to G. Allen, Sorex centralis 
Thomas, 191 1, Ann. Mag. N.H. 8: 758. Sayan Mountains, 100 miles west 
of Lake Baikal, 4,000 ft., Siberia.) Ranges to Mongolia. 

1913. Sorex baikalensis Ognev, Fauna Mosquensis, /.• 106. Zarentu Mountains, 


1914. Sorex araneus ultimus G. Allen, Proc. New England Zool. Club, j.- 51. Nijni 

Kolymsk, near mouth of Kolyma River, North-Eastern Siberia. 
1 91 4. Sorex macropygmaeus koreni G. Allen, loc. cit. 56. Nijni Kolymsk, near mouth of 

Kolyma River, North-Eastern Siberia. 
1921. Sorex macropygmaeus pleskei Ognev, Ann. Mus. Zool. Acad. St. Petersb. 22: 31 1. 

Charlamova Gora, Gdovsky district, Petrograd Govt., Russia. 
1 92 1. Sorex macropygmaeus rozanovi Ognev, loc. cit. 313. Listvenichnojc, west coast of 

Lake Baikal, Siberia. 
1921. Sorex macropygmaeus altaicus Ognev, loc. cit. 314. Ongudaj, Bijsk district, Tomsk 

Govt., Siberia. (Synonym, 1933, Sorex macropygmaeus altaicus tasicus Ognev, 

Abstr. Zool. Inst. Moscow Univ. /.' 62. Mouth of River Motliki, tributary 

of River Taza, Turukhansk district, Siberia.) 
192 1. Sorex amasari Ognev, Ann. Mus. Zool. Acad. St. Petersb. 22: 316. Valley of 

River Amazar, frontier between regions of Amur and Zabaikalje, Siberia. 
1921. Sorex ultimus petschorae Ognev, loc. cit. 317. Pvim-va, Petchora region, Northern 

1930. Sorex jenissejensis Dukclski Zool. Anz. 88: 77. Wostotschennje village, 40 

versts south-east of Minussinsk, Siberia. 
1933. Sorex ultimus midendorfii Ognev, Abstr. Zool. Inst. Moscow Univ. /.• 59. 

Sidorovsk, River Taza, Turukhansk district, North-\\'estern Siberia. 

(Synonym, 1933, Sorex ultimus middendorjii natio irkutensis Ognev, loc. cit. 60. 

Near Podunsk, on River Angara, Siberia.) 
1936. Sorex tundrensis europaeus Stroganov, Zool. J. Moscow, /j.- 130. Lake Chun, 

Imandra district. Kola Peninsula, North-Western Russia. 

Chinese and Japanese forms referred to macropygmaeus = caecutiens by Kuzyakin 
and Bobrinskii, or to buxtoni = caecutiens by G. Allen. 

1905. Sorex shinto Thomas, Abstr. P.Z.S. No. 23, 19. 1906, P.Z.S. igo^, 2: 338. 

Makado, Northern Hondo, Japan. 
1907. Sorex shinto saevus Thomas, P.Z.S. 408. Fifteen miles north-west of Korsakoff, 

Sakhalin Island. (Synonym, 1934, Sorex shinto savenus Tokuda, Zool. Mag. 

Tokyo, .}6: 578. ? misspelling oi saevus.) Occurs Hokkaido and Kurile Is. 
1907. Sorex annexus Thomas, P.Z.S. igo6: 859. Mingyong, iio miles south-east of 

Seoul, 1,300 ft., Korea. 




1QI2. Sorex cansiiliis Thomas, Ann. Mag. X.H. 10: 398. Forty-six miles south-cast of 
Taochou, Kansu, China. 

Miller (1912) treated the following form as a race of .S'. amneiis, but it seems to us 
to represent the present species. 

Sorex c.\ecutiens gr.w.vrius Miller, 1910 

I QIC. Sorex araneus granariiis Miller, .\nn. Mag. N.H. 6: 458. La Granja, Segovia, 
Since Miller published his Catalogue, the following form which is apparently 
referable to S. caccutiens has been named from Western Europe. 

1942. Sorex lapponieiis Melandcr, K. fysiogr. Sallsk. Lund. Forh. //.■ 134. Vittjarv, 
Northern Sweden. 

Sorex araneus Linnaeus, 1758 C:ommon Shrew 

Approximate distribution of species: Norway, Sweden, Britain, Channel Islands, 
France, Germany, Bohemia, Poland, Denmark, Holland, Belgium, Switzerland, 
Italy, Transylvania, Yugoslavia. Through much of the U.S.S.R., \vhcre the northern 
limit runs through the Eurasian tundra, and in many places reaches the Arctic coast; 
eastwards to the Pacific and Sakhalin; the southern limit skirts the steppes and semi- 
deserts of the northern Caucasus and Kazakstan, and the range includes Trans- 
caucasia, Mongolia, Manchuria. Bodenheimer lists the species from Palestine. Asia 
Minor. Apparently also frcxm Shensi, Kansu, Yunnan in China, and the Kurile 

Miller, 191 2, Cat. Mamm. IT. Europe, recognized the following eight races of this 
species in ^Vestern Europe. Some of them, however, are based on colour details 
which according to Kuzyakin and Bobrinskii are subject to wide individual variation 
and are likely to be useless fur diagnostic purposes. 

On the European forms see aho Zalesky, 1948, S.B. Osl. Akad. W'iss. 757.- 129. 

Sorex .\raneus ar.\neus Linnaeus, 1758 

1758. Sorex araneus Linnaeus, Syst. Nat. loth ed. /; 53. Upsala, Sweden. 
1828. Sorex coronatus Millet, Faune de Maine-et-Loire, /.• 18. Blou, Maine-et-Loirc, 

1828. Sorex perionatu\ Millet, loe. cit. (footnote). Not of Geolfroy, 1827. 

1829. Sorex dauheiitonii Q'.u\'\er,Kegn. Anim. i: 127. Not of Erxleben, 1777. 
1832. Sorex eoneinnus \\'agler, Isis, 2^: 54. Bavaria, Germany. 

1832. Sorex rhinolophui \Vaglcr, loe. eit. Bavaria. 

1832. Sorex melanodnn Wagler, loe. cit. Bavaria. 

1839. Sorex macrotriehu^ de Selys Longchamps, Etudes dc Micromamm. 20. No 

1839. .Sorex lahiosus ]cn\n^, .Vnn. N.H. 2: 326. Frankfurt, Germany. 
1847. Sorex vulficiris Nilsson, Ilium. Fig. Skand. Fauna, /.• 75. (teste Trouessart.) 
Range: Western Continental Europe, from Finland to France, Germany, Bohemia, 
Norway fpart). 



1780. Sorex tetragonurus Hermann, in Zimmerniann, Geogr. Gesch. 2: 383. Stras- 
bourg, Eastern France. 
1792. Sorex quadricaiidatus Kerr, Anim. Kingd. 208. Strasbourg, Eastern France. 

1834. Sorex hermanni Duvernoy, L'Institut, 299. 1835, Mem. Soc. Sci. Nat. Stras- 

bourg, 2: 3. Near Strasbourg, Eastern France. (Animal, not skull.) 

1835. Sorex fodiens Duvernoy, Mem. Soc. Sci. Nat. Strasbourg, 2: 17. (Skull, not 

animal.) Strasbourg, Eastern France. Not of Schreber, 1777. 
(?) i858. Sorex vulgaris pallidus Fitzinger, S.B. Akad. ^\'iss. \\'ien, ^j, 1 : 488. Locality 

unknown, probably Italy. 
i86g. Sorex vulgaris var. nuda Fatio, Faune Vert. Suisse, /.■ 127. Bernese Oberland. 
1869. Sorex vulgaris var. nigra Fatio, loc. cit. Lucerne, Switzerland. 

1900. Sorex vulgaris var. vel subsp. mollis Fatio, Rev. Suisse de Zool. 8: 471. Substi- 

tute for nigra. 

1901. Sorex araneus alticola Miller, Proc. Biol. Soc. Washington, /^.- 43. Meiringen, 

1905. Sorex vulgaris crassicaudatus Fatio, Arch. Sci. Phys. Nat. Geneve, ig, 4: 201. 

Zermatt, Switzerland. Not of Hemprich & Ehrenberg, 1834. 
1905. Crossopus ou Sorex ignotus Fatio, loc. cit. 202. (Mandible, not skull.) 
1905. Sorex araneus carpathicus Barrett-Hamilton, Ann. Mag. N.H. i^: 506. Hatszeg, 

Hunyad, 5,500 ft., Hungary. 
1927. Soricidus monsvairani Altobello, Rev. Franc. Mamm. /.• 6. Between Campo- 

basso and the Commune of Busso, Montevairano, Abruzzi, Central Italy. 

Status _^(3'e Guhno, 1939, Boll. Mus. Zool. Anat. Comp. Torino, 4j: 136. 
Range: Alps and neighbouring parts of Germany, France, Italy, east to Tyrol and 

Sorex araneus castaneus Jenyns, 1838 

1838. Sorex tetragonurus var. <^S{orex) castaneus ]fnyns, Ann. N.H. /.■ 424. Burwell Fen, 
Cambridgeshire, England. 

Sorex araneus euronotus Miller, 1901 

1901. Sorex araneus euronotus Miller, Proc. Biol. Soc. Washington, /^.- 44. Monlrejeau, 
Haute-Garonne, France. 

Sorex araneus santonus Mottaz, 1908 

1908. Sorex santonus Mottaz, Bull. Soc. Zool. Geneve, /.■ 118. Lignieres-Sonneville, 

Charente, France. 

Sorex araneus bergensis Miller, 1909 

1909. Sorex araneus bergensis Miller, Ann. Mag. N.H. 2- 4'6. Graven, Hardanger, 

Norway. Range: Western Norway, from Bergen region at least into 

Sorex araneus pyren.'^icus Miller, 1909 

1909. Sorex araneus pjrenaicus Miller, Ann. Mag. N.H. j.- 416. L'Hospitalet, Ariegc, 
4,700 ft., France. 



1009. Sorix arancusfn-tain Millcr, Ann. Mag. N.H. j: 415. Trinity, Jersey, Channel 


.Since Miller's C^atalogue (1912) the following names have been proposed for 
Western European forms of this species : 

iqr?. Sons araufits pfucimus Thomas, Ann. Mag. N.H. //.• 21G. Cliatal, Dobrudscha, 

Rumania. According to Ognev, ranges eastwards to Russia. 
if\\\. Sorcx uranlii Barrett-Hamilton & Hinton, Abstr. P.Z..S. 18. P.Z.S. 824. IsLmd 

of Islav, Hebrides. 
i()2(j. Sorex samniiicus A\U)hAh\ Boll. Inst. Zool. Rom.i, ;;.• 102. Pro\incc di C;ampo- 

basso, Goo-1,000 ni.. Southern Itah. 
Hr27. Soifx ariuiiin ehoiiintic W'ettstein, .\n/.. Akacl. W'iss. W'icii, i. Ruja, \'eliki lom 

\alley, south ol the .Mali Rainac, .Xcjrthern \'elebit, near Krasno, Croatia, 

l(|2o. Soiix aiaihi/s csikii G\ ula, .Mlatt. Ko/lcm Budapest, 2j: 54. 98. Mateszalka and 

Nagydobos Komitat, Szatmar district, Hungary. 
K130. Sorex araneiis holkavi Martino, Ann. H.N. Mus. Hung. 17.- 158. Igman Moun- 
tains, 1,350 m., Sarajevo district, Yugo.slavia. 
1037- ■^'''"•^' «""'"" pulchcr Zaleskv, Anz. Akad. Wiss. Wien, ^4: 214. Terscheling 

Island, HoUand. 
i<)3q. Sorex araniui pctrori M.irtino, Zap. Russk. Nauch. Inst. Byelgrad, 14: 90. Asan 

Cesma, Kozuii .Mountains, Southern Serbia, Yugoslavia. 
1944. Sorex araiiein bohemirus Stcpanek, Rozpr. Ceske Akad. Praha, 53, 2, No. 30: 2., Southern C'./,echoslo\akia. 

On account of individual variation, Kuzyakin and Bobrinskii think that the whole 
group of relati\elv large shrews in the U.S.S.R. should be classed as Sorex araneiis, 
without dixisioii into species and subspecies. Names available from the U.S.S.R. 
include .S'. daphaenodon (and races) which, in deference to Chaworth-Musters' opinion, 
is here listed as a distinct species. Besides these, the following, in order of naming, 
.ire available from the U.S.S.R.; 

1890. Sorex imilinenlatus Dobson, /\nn. Mag. N.H. -,: i l 5. Island of Sakhalin. Occurs 

1895. Sorex raddei Satuiiin, Arch. Naturgcsch, /.■ 109. Neighbourhood of Kutais, 

Georgia, Transcducasia (Ognev, 1938, Mamm. E. Europe, N. Asia, /.- 220). 

iSynonyms; .S'o;c.v />«//,( Thomas, 1913, .Xnn. Mag. N.H. //.■ 214. Sumela, 

30 miles south of Trebizond, 1,000-1,300 m., Asia Minor; and Sorex 

caiicasicus Satunin, 191 3, Trud. Obshch. Izuch. Chernomorsk. Poberezh. 2: 

24 i.V.I'.). Bakuryani, Tiflis Govt., Transcaucasia.) Clhaworth-Musters 

thought raddei was a distinct species. 
|i)o-,. Sorex araneiis horealis Kastschcnko, Rcc. Tomsk Univ. 83. Neighbourhood of 

Tomsk, Siberia. .According to G. .Allen, range includes mountain ranges 

Ironi Altai eastwards, and Mongolia. 
ii)i.;. ,SVj/,.v rohoratiii Hollister, Smiths. .Misc. Coll. Go, 24: 2. Ta]mcha, .Altai .Moun- 

t.iins, 12-, miles so\ith-east of Bijsk, Siberia. 
|i)i4 .So?, A I'll t;. Allen, I'roi . New England Zool. CUub, 5.- 52. Nijni Kolymsk, near 

mouth ol Kiplvm.i Ri\cr, .\oi th-Eastern Siberia. 


1914. Sorex asper Thomas, Ann. Mag. N.H. ij: 565. Tekes Valley, Tianshan Moun- 
tains, Central Asia. 
192 1. Sorex macroprgmaeus araneoides Ognev, Ann. Mus. Zool. Acad. St. Petersb. 22: 

315. Valley of River Sosnovka, east coast of Lake Baikal, Siberia. Bobrinskii 

and Kuzyakin consider this a form of 6'. araneus. 
1921. Sorex araneus tometuis Ognev, loc. cit. 329. Govt, of Tomsk, Siberia. 
1 92 1. Sorex araneus schnitnikovi Ognev, loc. cit. 330. Near Kopal, Semirechyia, East 

Russian Turkestan. 
1 92 1. Sorex araneus satunini Ognev, loc. cit. 331. Miusaret, Kars region, Caucasus. 
1 92 1. Sorex platycranius Ognev, loc. cit. 334. Near Nikolsk-Ussurijsky, Ussuri region, 

Eastern Siberia. 
192 1 . Sorex thomasi Ognev, loc. cit. 336. River Budarman, tributary of River Sosnovka, 

north-west coast of Lake Baikal, Siberia. 
1924. Sorex araneus tomensis isodon Turov, C.R. Acad. Sci. U.R.S.S. iii. River 

Sosovka, Bargusinsk taiga. Lake Baikal, Siberia. 
1928. Sorex araneus jaculensis Yinkchki, Zool Anz. y8: 102. Village of Sunlar, on the 

middle reach of the River Wiluj, Yakutsk, Siberia. 
193 1. Sorex vir turuchanensis Naumoff, Trans. Polar. Comm. Acad. Sci. U.S.S.R. .^.■ 

8-10 {N.V.). (See Ognev, Mamm. U.S.S.R. 2- 611.) Yanov Stan, River 

Turuchan, North-VVestern Siberia. 
(?) 1933. Sorex dukelskiae Ognev, Abstr. Zool. Inst. Moscow Univ. /.• 57. River 

Artyugin, tributary of Yenesei, Turukhansk district, Siberia. 
1933. Sorex araneus iochanseni Ognev, loc. cit. 61. Bobrovka, on River Kaba, in district 

of Altaiskaya, in former Semipalatinsk Province, Siberia. 
1933. Sorex araneus iiralensis Ognev, loc. cit. 62. Source of River Nyais, Northern Ural, 

Eastern Russia. 
1933. Sorex gravesi Goodwin, Amer. Mus. Novit. No. 637, i. Monoma River, 80 

miles east of Troitskov, Maritime Province, Eastern Siberia. 
1936. Sorex isodon ruthenus Stroganov, Zool. J. Moscow, /j.' 132, 141. Lake Seliger, 

Kalinin district, Russia. 

The following Chinese names are likely to represent .S'. araneus: 
Sorex araneus sinalis Thomas, 191 2 

1912. Sorex sinalis Thomas, Ann. Mag. N.H. 10: 398. Forty-five miles south-east of 
Feng-hsiang-fu, Shensi, 10,500 ft., China. Ranges to Southern Kansu. 

Sorex (?) .\raneus excelsus G. Allen, 1923 

1923. Sorex excelsus G. Allen, Amer. Mus. Novit. No. 100, 4. Summit of Hoshan, 

Peitai, 30 miles south of Chungtien, Yunnan, 13,000 ft., China. 
The following form from Japanese territory is likely to represent .S". araneus: 
1933. Sorex megalotis Kuroda, Bull. Biogeogr. Soc. Jap, 4, i; 47. Chikuradake, 

Paramushiru, Kurile Islands. Synonym: 
(?) 1930. Sorex paramushirensis Kishida, Z. Mag. Tokyo, XLII, 373, nom. nud. 

Sorex daphaenodon Thomas, 1907 

Approximate distribution of species: Sakhalin, Kurile Islands, Central and Eastern 
Siberia, Hokkaido in Japan. Referable to S. araneus according to Kuzyakin and 


PALAEARC:TR: A\U INDIAX mammals ly-.S I94f> 

1907. Soii'x dajihaenudon Thomas, P.Z.S. 407. Darinc, 25 miles north west of Korsa- 

kofl, .Sakhalin Island. The following forms were referred to the synonymy 

of this in Chaworth-Musters' M.S. 
IQ14. Sorex sangidnidcns G. Allen, Proc. New England Zool. Club, 5.- 54. Xijni 

Kolymsk, near mouth of Kolyma River, North-Eastern Siberia. 
192 1. Sorcx iibiriemis Ogncv, Ann. Mus. Zool. Acad. St. Pctersb. :?:?.• 328. \'illage 

Koltchugina, Kuznetsky district. Govt, of Tomsk, Siberia. 
1924. Sorcx daphaenodon yeiocmis Kishida, Monogr. Jap. Mamm. 168. Kuroda, 1928, 

J. Mamm. g: 222. Province of Ncmuro, Hokkaido, Japan. 
1933. Sortx daphaenodon orii Kuroda, Bull. Biogeogr. Sue. Japan, ^, i : 48. Nasauki, in 

Paramushiru, North Kurile Islands. ( Sonw mil Kishid.i, 1930, Zool. Mag. 

Tokyo, ^3: 373, nom. mtd.) 
1933. Sorcx daphaenodon icaloniOgnev , Abstr. Zool. Inst. Moscow Univ. /.' 63. Mouth 

of River Motliki, basin of River Taza, Turukhansk district. Northern 

Central .Siberia. 

Sorex buchariensis Ogne\ , 1921 

Approximate distribution of species: Russian Pamir Mountains. 


1 92 1. Sorex buchariensis Ognev, Ann. ,\Ius. Zool. .\cad. St. Petersb. 22: 320. \'alley of 
River Da\an-su, North- West Russian Pamir Mountains. 

Sorex pacificus C^oucs, 1877 Giant Shrew 

Approximate distribution of species: Ussuri region of Eastern Siberia; North- 
Western United .States fCaiifornia, Oregon). For note on status of Ussiui form, see 
above (page 441. 

Sorex pacificus Coues, 1877 

1877. Sorex pacificus Clones, Bull. U.S. Geol. & Geogr. Surv. Terr, jj.' 650 (jV.F.). 

Fort Umpqua, mouth of Unipqua River, Douglas County, Oregon, U..S.A. 
(?) 1937. Sorex mirahiUs Ot,'nev, Bull. Soc. Nat. Moscou, Sect. Biol. 46: 268, 270. 

Kiskinka Ri\cr willey, Ussuri region. Eastern Siberia. 

Sorex alpinus Schinz, 1837 .Alpine Shrew 

.\]3proximate distribution of species: France, south to Pyrenees, Germany, 
S\sitzerland, Italy, ^'ugoshuia, Transylvania, Prjland. 


1837. Sorcx alfiinus Schinz, Ncue Denkschr. Allgem, .Schweiz. Gesell. Naturwiss. 

Neuchatel, /.• 13. St. Gothard Pass, .Switzerland. 
(?j 1840. Sorex antinorii Bonaparte, Iconogr. Faun. Ital. /.■ fasc. 29, no exact locality, 

"probably not a European species" (Miller, 1912). 
(?) 1870. Sorcx uilcrmedius C^ornaiia, Catal. Descr. Mamm. Ital. 27. Hills of Brianza, 

Como, Italy. (Part, body. See Sordelli, i8f)i).) 



(?) 1899. Sorex alpinus var. longobarda SordelLi, Atti Soc. Ital. Sci. Nat. Milano, ^8: 

363. (Synonym ot interrnedius.) 
Range: France, Switzerland, Transylvania (Pyrenees, Jura, Alps, Tyrol, etc.). 

Sorex alpixus hercynicus Miller, 1909 

1909. Sorex alpinus hercynicus Miller, Ann. Mag. N.H. j.- 417. Mauseklippe, Bode 

Valley, Harz \Iountains, Germany. Range : Harz Mountains and Riesenge- 

birge, Germany. 

Sorex cylindricauda Milne-Edwards, 1872 Stripe-backed Shrew 

Appro.ximate distribution of species: Yunnan, Szechuan, Kansu, Shensi, in China; 
and Northern Burma. 

Sorex cylindricauda cylindricauda Milne-Edwards, 1872 

1872. Sorex cylindricauda Milne-Edwards, Xouv. Arch. Mus. H.X. Paris, 7. Bull.: 92 

(footnote). Moupin, Western Szechuan, China. 
191 1. Sorex bedfordiae Thomas, Abstr. P.Z.S. No. 90, 3. P.Z.S. 164. Omisan, 9,500 ft., 

Szechuan, China. 
191 1. Sorex wardi fumeolus Thomas, Abstr. P.Z.S. No. 100, 49. 1912, P.Z.S. 132. 

\Veichoe, on Siho River, \Vestern Szechuan, 6,000-11,000 ft., China. 

Sorex cylindricaud.-^ wardi Thomas, 1 9 11 

igii. Sorex wardi Thomas, Abstr. P.Z.S. No. 90, 3. P.Z.S. 165. Taochou, 9,000 ft., 
Kansu, China. 

Sorex cylindricauda gomphus G. Allen, 1923 

1923. Sorex bedfordiae gomphus G. Allen, Amer. Mus. Novit. No. 100: 3. Mucheng, 

Salween drainage, \Vestern Yunnan, 7,000 ft., China. Ranges to Northern 


Incertae sedis 

Sorex pusillus Gmelin, 1774, Reise, 5.' 499, pi. 57, fig i. Persia, no exact localit\-. 
Perhaps a Crocidura, and probably unidentifiable. 

Sorex (?) shinanensis (described as Urotrichus talpoides shinanensis) Yagi, 1927, Zool. 
Mag. Tokyo, jg: 201 (A'.IVi. Kitazawatoge, between Senjogatake and 
Higashi-Komagatake, in Southern Japanese Alps, Hondo, Japan. Status 
Jide Kuroda, 1938, List Jap. Mammals. Synonym, possibly, 1937, Sorex 
dorichurus Kishida, Rigaku Kai, 25, ^o. 410: 742. Senjogatake, Southern 
Japanese -Alps, Hondo. [N.V.) 

Genus BLARINELLA Thomas, 191 1 
igii. Blarinella Thomas, P.Z.S. 166. Sorex quadraticauda Milne-Edwards. 
I species: Blarinella quadraticauda, page 56 



Blarinella quadraticauda Milnc-Kdw aids, ifjyj Short-taiied Moupin Shrew 

Appniximatc distribution ol species: Szcchuan, ^'ullnan anil Kansn, China; 
Niirthern Burma. 


1872. .S'o;v.v (juadraliiduda Milne-Edwards. Reeh. H.X. Manim. 261, pj. 38a, furs. 
■:;--!d. pi. 38b, fi<4. 2. .Moupin, Szcrhuan. C^hina. 


101-'. I'llariritlla c^risthla Thmnas, .Ann. .Mat;. X.H. lo: 4110. FortN-two miles south- 
cast of Taocluiii. 10,000 It., Kansu, Cihina. 

BL.\RINt.LL.\ qU.XDRATICAt-'D.V WARD! ThoHias, I f ) I 5 

It)!;,. Blarinella ky;r/; Thomas, .\nn. Mas;. X.H. i§: 336. Pipimaw, 26 X., 08.3'/ E.. 
8,000 ft.. Upper Burma. Range includes Yunnan. 

Genus SORICULUS Blvtii, 185.1 

1854. Sdiiciiltii Bhth, J. .'Vsiat. Soc. Bengal, 2;]: 733. (,'orsira ni<^re\criii Gray. 
1907. (.hodsigoa Kastschcnko, Ann. .\Ius. Zonl. Arad. .St. Petersb., 10: 251. Soriculus 
salrnskii Kastschcnko see G. Allen, 1038: 1041. \'alid as a subgenus. 
E/iiuirniiliis iiihgen. nor. Type species: .S'o/v.v laiidalus Horsfield. \'ali(l as a 
subgenus, to contain also .S'. hucops Horsfield.' 

6 species: Soricnlus caiidatus, page '>') 
Soriculus kvpsibius, page bo 
Soriculus leucops, page j(\ 
Soriculus lowci, page ')i 
Soricnlus ni,i;resccns. page -,8 
Soriculus salcnskii. page <>o 

( )1 these species, h\]>\ihius, salcnskii and lohti belong to the subgenus C/iodsii^oa 
which 28 instead of 30 teeth. The vanishing tooih the last upper unicuspid) is 
\estigial in the other species and, as has already been poiiit<-d out by Osgood, its 
presence can at most be of subgencric value. The first nam ■ 111 the subgenus Chndsigoa 
is .S. lirpsihiu^. a relati\eh- short-tailed species in which the liindl(iol is about 
ii-i"! mm., and according tn G. .Mien it occurs in parts nf China with an allied 
longer-tailed s]3ecies hitherio known as unillii, but here (diisidered as representing 
the earlier name salcnskii. This has the hincHoot about i(i-2ii mm. Typical salcnskii 

' Subgenus Chodsiguir, with ci'^ht ui)prr t'-cth. 

Subgenus Smiculns: with nine upper teclh. the \ery small extr.i upp.r unicuspid being present. 
F'; tail short, usually less than 70 per cent, o! liead and body lin all bul radulm averages less 
than Ijo per cent.;. Rather large, head and body usually more than 70 mm. Foreclaws enlarged. 

Subgenus K/mni Ifidin; with nine upper teeth, the very small extra upper unicuspid being present. 
Not fussorial; tail long, approximatclv 110-147 piT rent, avi-rage of hear! and bodv. Small, head and 
body length usually less than 70 mm. ■ prrliaps excepting S. c. bailryi . Forn l.n\> not enlarged. 



seems to be only known by one specimen, which has an unusually long tail (over 
140 per cent, of head and body length) and a hindfoot of 20.5 mm., but according to 
Anthony, 1941, Field Mus. Publ. ^ooL 2j: 71, the hindfoot in forms which he 
referred to smithii can be as much as 20 mm. ; and as in some forms currently referred 
to smithi the tail is also considerably longer than the head and body (though less 
elongated than in the type oi salenskii), there seems not much reason why the name 
salenskii should not be regarded as the prior name for the smithi section of races. The 
third species of Chodsigoa, lowei, has a short hindfoot, as in hypsibius, but a very long 
tail, as is often the case in salenskii, combined with some cranial peculiarities pointed 
out by its describer, and although not well known is tentatively regarded as valid. 
In those species hitherto referred to Soriculus (with 30 teeth), there are two very 
distinct groups. The type, nigresceris, is a rather large, heavily built fossorial shrew 
with enlarged foreclaws and a short tail which is rarely as much as 70 per cent, of 
the head and body. The other two species, caudatus and leucops are rather small, 
slender shrews with small foreclaws and a long tail which is on average go per cent. 
or more of the head and body fpossibly excepting the very little-known Formosan 
race) (the species caudatus as a rule has the tail 90-109 per cent, of the head and body, 
and the species leucops has it nearly half as long again as the head and body) . The 
external difference between the nigresceris group and the caudatus-leucops group is so 
well marked that we feel subgeneric division is advisable, and propose the name 
Episoriculus, with type S. caudatus. The distinction between the t-\vo subgenera is 
greater than between Sorex and Blannella in external characters. It is necessary to 
note that Blanford used the name S. macrurus for S. leucops, but macrurus was a nomen 
nudum except from Blanford (1888), and Osgood has shown that the name leucops has 
priority dating from 1855. G. Allen regarded the form sacratus as a race of 5. caudatus, 
but more recently Anthony (1941) has revived sacratus as a specific name because a 
form which he calls a race of sacratus occurs with caudatus in Northern Burma. He 
suggests (page 69) that it is possible that the two animals have different habitat 
preferences and thus remain separated in the same locality; until the contrary is 
proved we follow G. Allen. Two rather differentiated forms, radulus and batleji, ha\-e 
been named since Blanford classified the Indian species, but neither are so distinct in 
our opinion that they need be given specific rank; the first is here considered to 
represent nigresceris, and baileji is here referred to caudatus. The status of the t >vo 
named Formosan forms of the genus is not clear. The pigmentation on their teeth is 
extremely weak or untraceable, and the Formosan Chodsigoa is based on a skull, the 
skin being unknown. 

The available species of Soriculus may be distinguished as below: 

I. Eight upper teeth. Foreclaws not enlarged. Subgenus Chodsigoa.) 2 

Nine upper teeth, the last upper unicuspid exceedingly reduced. 4 

2. Hindfoot about 16-20 mm. (Tail usually as long as or longer than head and 
body.) Soriculus salenskii 

(Form seen: smithi.) 

Hindfoot about 11-15 mm. 3 



■3. Tail much lunger than head and body. Soriculiis lowei 

(Type in B.M.) 
Tail about 80 per cent., or less, of head and body. Soriculiis hypsibius 

(Forms seen: hvpsihius, larvariim. lamia.) 

4. Tail normally less than three-quarters of head and body length, its length 50 mm. 

and less. Foreclaws enlarged. (Subgenus Soriculiis.) Soriculus nigrescens 
(Forms seen: nigrescetn, centralis, caurinus, pahari, radulus.) 
Tail normally 90 per cent, or more of head and body, its length usually over 
-,o mm. Foreclaws small. (Subgenus Episoriculus.) 5 

5. Tail about 90-109 per cent, of head and body, its length below 80 mm. 

Soricului caudatus 
(Forms seen: caudatus, sacratus,Jumidus (no measured skins), bailcvi.) 
Tail as a rule about 1.1.5 P*^'' cent, of head and body, its length usually over 80 mm. 

Soriculus Icucops 
(Forms seen : Icucops, macrurus, Irene.) 

Subgenus SORICULUS BIyth, 1854 

Soriculus nigrescens Gray, 1842 Sikkim Large-clawed Shrew 

Appri.>ximatc distribution of species: Bhutan, Kumaon, Sikkim, Nepal, Mishmi, 
Northern Burma. 

Soriculus nigrescens nigrescens Gray, 184? 

1842. Corura nigrescens Gray, .\nn. -Mag. X.H. 10: 261. Darjecling, India. (Hinton, 

1842. Sorcx aterrinius Blyth, J. Asiat. Soc. Bengal, 12: 928, nom. nuil. 1854, J. Asiat. 

Soc. Bengal, I'j: 733. Darjceling. 
1849. Sorcx sikimensis Hodgson, Ann. Mag.- N.H. 5.- 203, nom. nud. 1855, J. Asiat. 

Soc. Bengal, 16: iii. Darjceling. 
18G3. Sorex oligurus Gray, Ciat. H(jdgson C<A\. .\epal & Tibet, 2nd cd. 8, Sikkim 

nom. nud.). 
18(^3. Sorex holosericeus Gray. loe. cit. 9. Darjecling ^nom. nud.). 

Soriculus nigrescens radulus Thomas, 1922 

ii|22. Soricului radulus Thomas, J. Bombay N.H. Soc. 28: 429. Dreyi, 5,140 ft., 
.Mishmi Hills, Northern Assam. Ranges to North Burma. 

Soriculus nigrescens t.vhari Hinton, 1922 

1922. Soriculus nigrescens pahari Hinton, J. Bombay N.H. Soc. 28: 1053. Gnatong, 
12,300 ft., Sikkim. 

Soriculus nigresce.vs C-\urixus Hinton, 1922 

i')22. Soriculus nigrescens caurinus Hinton, J. Bombay .\.H. Soc. 28: 1054. Khati, 
7,l)0o it., Kumaon, Northern India. 



1922. Soriculus nigrescens centralis Hinton, J. Bombay N.H. Soc. 28: 1054. Bouzini, 


Subgenus EPISORICULUS Ellerman & Morrison-Scott, 1951 

Soriculus caudatus Horsfield, 1851 Hodgson's Brown-toothed Shrew 

Approximate distribution of species: Kumaon, Sikkim, Mishmi, Northern Burma; 
Szechuan, Yunnan and apparently Formosa; Tonkin, in Indo-China. 

Soriculus caudatus caudatus Horsfield, 1851 

1 85 1. Sorex caudatus Horsfield, Cat. Mamm. Mus. E. India Co. 135. Darjeeling, 

India {fide Chaworth-?>Iusters). 
(?) 1863. Sorex homourus Gray, Cat. Hodgson's CoU. B.M., 2nd ed. 8, nom. nud. Sikkim. 
1877. Soriculus gracilicauda Anderson, J. Asiat. Soc. Bengal, 46, 2: 282. Sikkim. 
1890. Soriculus minor Dobson, Monogr. Insectiv. jj, pi. xxiv, figs. 2-2b. Manipur. 
Range: Kumaon, Sikkim, Northern Burma. 

Soriculus caudatus sacratus Thomas, 1 9 11 

191 1. Soriculus sacratus Thomas, Abstr. P.Z.S. 4. P.Z.S. 165. Omei Shan, 6,000 ft., 
Szechuan, China. 

Soriculus (?) caudatus fumidus Thomas, 19 13 

1913. Soriculus fumidus Thomas, Ann. Mag. N.H. //.• 216. Mt. Arizan, 8,000 ft., 

Central Formosa. 

Soriculus caudatus baileyi Thomas, 1914 

1 9 14. Soriculus baileyi Thomas, J. Bombay N.H. Soc. 22: 683. Tsu River, Mishmi 

Hills, 7,500 ft., north of Assam. Range includes Tonkin, Indo-China. 

Soriculus caud.jltus umbrixus G. Allen, 1923 

1923. Soriculus caudatus umbrinus G. Allen, Amer. Mus. Novit. No. 100, 5. Mucheng, 

Salween drainage. South- Western Yunnan, 7,000 ft., China. Ranges into 
Northern Burma. 

Soriculus leucops Horsfield, 1855 Indian Long-tailed Shrew 

Approximate distribution of species: Nepal, Sikkim, Northern Burma; Szechuan 
and Yunnan, China. 

Soriculus leucops Horsfield, 1855 

1855. Sorex leucops Horsfield, Ann. Mag. N.H. 16: iii. Nepal. 

(?) 1863. Sorex nivicola Gray, Cat. Hodgson's Coll. B.M., 2nd ed. 8, nom. nud. 

1863. Sorex macntrus Hodgson, loc. cit. 9, nom. nud. Not macrourus Lehmann, 1822. 

1888. Soriculus macrurus Blanford, Fauna Brit. India, Mamm. /.■ 231. Darjeeling, 

Northern India. 
191 1. Soriculus Irene Thomas, Abstr. P.Z.S. 49. 1912, P.Z.S. 132. Yuanchinghsien, 

South-Western Szechuan, 5,200 ft., China. 

E 59 


Subgenus CHODSIGOA Kaststhcnko, 1907 

Soriculus hypsibius dc Winton, 1899 dc \Vinton's Shrew 

Approximate distribution of species: Yunnan and Szim huan, northwards to 
Kansu, Shcnsi and ClhihH, China. 

SoRiciLis iivpsinn s iivpsiBUs dc Wintim, iflf)i) 

1899. Soriculu.s hv/iMhiiis dc \\'inton, P.Z.S. 574. \'ant;liupa, North-Western Szcchuan, 

1907. .Soriculus [Chod^igoa) benxowikii Kastschcni<o, Ann. Mus. Zool. Acad. St. 

Pctcrsb. in: 252. Chodsigou, Northern Szcchuan. 
Range : Yunnan (part), Szcchuan, Shcnsi. 

Soriculus hypsidrls larv.xrum Thomas, 191 1 

1911. Chodsiaoa Inrvaium Thomas, Abstr. P.Z.S. 49. 1912, P.Z.S. 133. Eastern Tombs, 

()5 miles east of Pekin, 1,000 it., Clhilili, Ciiina. 

Soriculus hypsibius lamula Thomas, 19 12 

191 2. Chodsiona lamula Thomas, Ann. Mag. N.H. 10: 399. Forty-six miles south-east 

of Taochou, Kansu, 9,500 ft., C^hina. 


1923. Chodsiiioa hv/iiibia parva G. .Allen, Amcr. Mus. Novit. No. loo: 5. Ssushanchang, 
Likiang Range, Western Yunnan, 9,000 ft., China. 

Soriculus salenskii Kastschenko, 1907 Salenski's Shrew 

Approximate distribution of species: as here understood, Shcnsi, Szechuan and 
Yunnan, in (Ihina; Northern Burma. 

Soriculus salen.skii salenskii Kastschenko, 1907 

1907. Snriciiliis iChndiigoa) salenskii Kastschenko, Ann. Mus. Zool. Acad. Sci. St. 

Pcteisb. 10: 253. G. Allen, 1938, Mamm. China & Mongolia, /.• 108. 

Linganfu, Northern Szechuan, China. (Tate (1947) thinks this is a distinct 

species from .S'. smithii on account of its longer tail.) 

Soriculus sale.nskii smithi Thomas, 191 1 

191 1. Clwdsiooa smithii Thomas, Abstr. P.Z.S. 4. P.Z.S. 166. Tatsienlu, Szechuan, 
9,000 It., China. Range includes Tsingling Mountains, Shcnsi, China. 

SoRir:ULUS SALENSKII PARC,\ G. Alicu, 1 923 

1923. Clindsii^oa smilhii parca G. Allen, .'\mer. Mus. Novit. No. loo: 6. Homushu 
Pass, Western Yunnan, 8,000 ft.. C;hina. Ranges to Northern Burma (part). 

Soriculus salenskii eik\'a Anthony, 1941 

1941. Chodsinoa smithii furva Anthony, Field Mus. Publ. Zool. 2j: Ji. .Mt. Iniaw Bum, 
I), (Mill It., \i)rtli(rn l?urma. 


Soriculus lowei Osgood, 1932 Lowe's Shrew 

Approximate distribution of species: Tonkin, in Indo-China. 

Soriculus lowei Osgood, 1932 

1932. Chodsigoa lowei Osgood, Field Mus. Publ. Zool. 18: 249. Chapa, Tonkin, 

Incertae sedis 

1913. Chodsigoa sodaiis Thomas, Ann. Mag. N.H. //.• 217. Mt. Arizan, 8,000 ft., 
Central Formosa. Based on a single skull with scarcely pigmented teeth; 
skin unknown. 

Genus NEOMYS Kaup, 1829 

1829. JVeomys Kaup, Skizz. Europ. Thierwelt, /.• 117. Sorex daubentonii Erxleben = 

Sorex fodiens Pennant. 
1829. Leucorrhynchus Kaup, loc. cit. 118. Sorex lineatus Geoffro)' = Sorex fodiens 

1829. Hydrogale Kaup, loc. cit. 123. Sorex remijer GcofTroy = Sorex fodiens Pennant. 
1832. Crossopus Wagler, Isis, 275. Sorex fodiens Pennant. 
1835. Hydrosorex Duvernoy, Mem. Soc. Sci. Nat. Strasbourg, 2: 19. Sorex fodiens 

1835. Amphisorex Duvernoy, loc. cit. 23. Sorex herinanni Duvernoy = Neomys fodiens 

skull plus Sorex araneus tetragonnriis, skin. 
1838. Pinalia Gray, P.Z.S. iS^j: 126. Synonym oi Crossopus ex Gray M.S. 

2 species: Neomys anomaltis, page 64 
Neomys fodiens, page 61 

This genus was dealt with at some length by Miller, 191 2, Cat. Mamm. W. Europe, 
65. Bobrinskii recognizes two species only, which are both compared in Miller (who 
subdivided anomalus). 

Neomys fodiens Pennant, 1771 European Water-Shrew 

Approximate distribution of species: Britain, France (south to Pyrenees), Denmark, 
Belgium, Holland, Switzerland, Italy, Transylvania, Germany, Norway, Sweden, 
Finland ; in Russia the northern limit runs almost along the coast of the Arctic 
Ocean, and in Western Siberia a little south of the Arctic Circle (apparently to 
about Lake Baikal) ; in the Far East there have been individual finds on the lower 
Amur and coast of Sea of Okhotsk, and Sakhalin. The southern limit skirts the 
Northern Caucasus, the Volgo-Ural and Kazakstan steppes. Bodenheimer recorded 
this species from Palestine. But it seems more likely that the Palestine form is 
anomalus, since the latter is the water-shrew of Asia Minor and the Mediterranean 



Neomys fodiens fodiens Pennant, 1771 
771. Sorex fodiens Pennant, Synopsis Qiiadrupcds, 308. Berlin, Germany. 
{Sorex fodiens Schrcbcr, 1777, Saugcth, 7; 57 i. Berlin, Germany.) 

776. Sorex aqiialieiis Mullcr, Natursyst. Suppl. u. Resist. Band. 36. France. Not of 

Linnaeus, 1758. 

777. Sorex dauhentomi Erxleben, Syst. Rcsn. Anim. /.• 124. Burgundy, France. 
780. Sorex earinatus Hermann, in Zimmcrmann, Geogr. Gesch. 2: 383. Strasbourg, 

Eastern France. 
79J. Sorex liricaudatus Kerr, Anim. Kingd. 208. Strasbourg, Eastern France. 
793. Sorex fluviatilis Bechstein, Gemeinn. Nat. Deutschlands, j.- 746. (Suggested, 

but not adopted, as preferable to fodiens.) 
793. Sorex eremita Meyer, Zool. Annalen, /.• 323. Thuringia, Germany. 
?) 1800. Sorex eanieulariiis Bechstein, Thomas Pennant's AUgcm. Uebers. Vierf. 

Thiere, 2: 541. Renaming oi fodiens Bechstein, 1793. 
800. Sorex fodiens alhus Bechstein, loc. cil. 723. 

811. Sorex hydrophilus Pallas, Zoogr. Rosso. Asiat. 130. Berlin, Germany. 
811. Sorex lineatus Geofi'roy, Ann. Mus. H.N. Paris, ij: 181. Paris, France. 
811. Sorex remifir Geofiroy, loc. cit. 182. Abbeville, Somme, France. 
818. Sorex eoUaris Desmarest, Nouv. Diet. H.N. 22: 65. Islands at mouth of Escaut 

and Mcusc, Holland. 
822. Sorex maerourus Lchmann, Obs. Zool. Faun. Hamburg, /.• 5. Sachsenwald, 

Schleswig-Holstcin, Germany. [N.V.) 
826. Sorex ampliibiiis Brehm, Ornis, :?.• 38. Renthendorf, Thuringia, Germany. 
826. Sorex natans Brehm, loc. cit. 44. Renthendorf, Thuringia, Germany. 
826. Sorex stagnatUis Brehm, loc. at. 47. Renthendorf, Thuringia, Germany. 
830. .S'orc.v TOfl/w Brehm, Isis, 1128. Renthendorf, Thuringia, Germany. 
832. Sorex miisculiis \Vagler, Isis, 54. Bavaria, Germany. 
832. Sorex psilurus VVagler, loc. cit. Bavaria, Germany. 

834. Sorex nioripes Melchior, Den Danskc Stats og Norges Pattedyr, 68. Sielland, 


835. Sorex hermanni Duxcrnoy, Mem. .Soc. Sci. Nat. Strasbourg, 2: 23. (Part; the 

skull only; the skin is another form.) Strasbourg, Eastern France. 
838. Amphisorex linneana Gray, Ann. N.H. 2: 287. North Bothnia, Sweden. 

838. Amphisorex eonstrictiis Duvernoy, Mem. Mus. H.N. Strasbourg, Suppl. 2: 4. 

839. Sorex fodiens var. Uucotis de Selys Longchamps, Etudes de Micromamm. 142, 

norn. niid. 
839. Sorex fodiens var. alhiventris de Selys Longchamps, loc. cit., nam. nud. 
?) 1845. Sorex fodiens nigricans Nilsson, Atti deUa scsta Riunione degli Sci. Ital. 

Torino, 1844: 357. Sweden {nom. nud.). 
868. Sorex fonhriatiis Fitzingcr, S.B. Akad. \Viss. \\'ien. 57, i : 610. Not of Wagler, 

868. (irossopus cihatus griseogiilaris Fitzinger, loc. eit. 623. Chartres, Eure-et-Loire, 

870. Sorex intermedins Gornalia, Cat. Desc. Mamm. Ital. 27. Hills of Brianza, Como, 

Italy. iPcUt, tail only. See Sordelli, 1899.) 
899. Sorex alpimis var. longobarda Sordelli, Atti Soc. Ital. Sci. Nat. Milano, 38: 363. 

MS. synonym of intermedins. 
901. Neomys fodiens minor Miller, Proc. Biol. Soc. Washington, 14: 45. Montrejeau, 

Hautc-Gari inne, Fr.uice. 


1905. Crossopus ou Sorex ignolus Fatio, Arch. Sci. Phys. Nat. Geneve, ig, 4: 202. 

Switzerland. (Skull, not mandible.) 
1905. jYeomjs fodiens naias Barrett-Hamilton, Ann. Mag. X.H. /j.- 507. Hatszeg, 

Hunyad, Hungary. 
igo6. Neomys fodiens nanus Lydekker, Zool. Record, ^2, Mamm. 34. Accidental re- 
naming oi naias. 
(?) 1 914. Neomjs leptodactylus Satunin, Mitt. Kauk. Mus. 8: 90. Kasikoporan, 

(?) 1924. Neomys fodiens alpestris Burg, \Veidmann, Pallasia, 2, 2: 90. Engadine {nom. 

nud. Original N.V.). 
1926. jNeomys fodiens halkaricus Ognev, Bull. Sci. Inst. Explor. Caucasus, /.• 42, 55. 

Neighbourhood of the town of Nalchik, Terek region, Caucasus. 
1 93 1. Neomys fodiens slresemanni Stein, Mitt. Zool Mus. BerUn, ly: 278. (Status _^f/(' 

Pohle, 1933.) Reipzig, near Frankfurt-on-Oder, Germany. 
Range: Norway, Sweden, Belgium, France, Germany, Hungary, Switzerland, Italy, 
to Russia, Transcaucasia and Western Siberia. 

Neomys fodiens bicolor Shaw, 1791 

1 791. Sorex bicolor Shaw, Naturalist's Miscell. 2, pi. 55. Oxford, England. 

1805. Sorex ciliatus Sowcrby, Brit. Misc. ^g: 103. Norfolk, England. 

1838. Amphisorex pennant a Gray, P.Z.S. iSjj: 125. England. 

1840. Crossopus sowerbyi Bonaparte, Iconogr. Faun. Ital. /, fasc. 29, in text under 

C. fodiens. 
Range: England, \Vales, Scotland. 

Neomys fodiens orientis Thomas, 191 4 

1914. Meomys fodiens orientis Thomas, Ann. Mag. N.H. ij: 564. Swamps of River 

Kammanajaretschka, near Djarkent, Semirechyia, Russian Central Asia. 

1915. Meomys fodiens orientalis Hinton, Zool. Record, 5/, Mamm. (1914) 44. Acci- 

dental renaming of orientis Thomas. 
(?) 1 92 1. Neomvs fodiens brachyotus Ognev, Ann. Mus. Zool. Acad. St. Petersb. 22: 343. 

Near Kopal, Semirechyia, Russian Central Asia. 
(?) 1 92 1. Neomys argenteus Ognev, loc. cit. 346. Coast of Lake Baikal, Siberia. 
Range: Bobrinskii quotes brachyotus from Semirechyia, the Altai, Tarbagatai Moun- 
tains, Central Siberia and the Far East, but orientis antedates. 

Neomys fodiens dagestanicus Heptner & Formozov, 1928 

1928. Neomys fodiens dagestanicus Heptner & Formozov, Zool. Anz. yy: 273. Fort 
Gunib, 6,000 ft., Daghestan, Eastern Caucasus. 

Neomys fodiens watasei Kuroda, 1941 

1941. Neomys fodiens watasei Kuroda, Bull. Biogeogr. Soc. Tokyo, //.• 114. Toyohara 

City, Sakhalin Island. (Neomys watasei Kishida, 1930, Zool. Mag. Tokyo, 

42: ^■j2, nom. nud.). 
Incertae sedis 

1913. Neomys schelkovnikovi Satunin, Trud. Obshch. Izuch. Chernomorsk. Poberezh, 
2: 24. [N.V.) Ushkul village, Svanetiya, Transcaucasia. (Chnworth-Musters 
regarded this as a form oi N. fodiens.) 



Neomys anomalus Clabiern, 1907 Mediterranean \\'atcr-Shrevv 

Approximate distribution ot'species: Spain, Switzerland, Italy, Carpathian Moun- 
tains, Pyrenean France, Yugoslavia, Greece, Poland, Crimea, Ukraine to Voronezh 
rcsjion in Russia, and Asia Minor. 

Xeomys anomalus anomalus Cabrera, 1907 

1907. .Xeomys anomalus Cabrera, Ann. Mag. N'.H. ^o: 214. i September 1907. San 
Martin de la \'ega, Madrid, Spain. 

Xeomys .\nom.\lus milleri Mottaz, 1907 

1907. .Xcoinys milhri Mottaz, Mem. Soc. Zool. France, 20: 22. 20 September 1907. 

Cliesieres, Alpes \'.iudoises, 1,230 m., Switzerland. 
192 1 . Xfornvs sorkioides Ognc\ , .Ann. Mus. Zool. .\cad. St. Pctersb. -^:?; 347. Beloviczh, 

Grodno district, Poland. 

Xeo.mys .\nom.\lus teres Miller, 1908 

H)o8. .Womrs /(7('-v Miller, .Ann. Mag. X.H. /.• 68. TwentN-five miles north of 
Erzerum, 7,000 It., Asia Minor. 

Xeomys .^nom-^lus mokrzeckii Martino, 19 17 

191 7. Neomys fodicns mokrzeckii Martino, Bull. Soc. Xat. Crimee, y: i (of reprint). 
Kholodnaya \Vater, River Alma, Crimea. ^.Although this form was named 
as a race offodiens, Bobrinskii states that that species is absent from Crimea, 
and that only .A . anomalus occurs there.) 

Xeomys ano.m.^lus josti Martino, 1940 

1940. .Xeomys milleri josli Martino, Ann. Mag. X.H. j.- 494. Ohrid, Macedonia, 
Southern Vugosla\ia. 

Genus SUNCUS fLhrcnberg, 1833 

1833. Siinais Ehrenberg, in Hemprirh & Ehrenberg, Symb. Phys. Mamm. 2: k. 

Suncus sacer Ehrenberg. 
1839. Pachyura de Selys Longchamps, Etudes de .Micromamm. 32. Sorex etniseiis 

1843. .SwH/tf/i Sundevall, K. Svcnska. \'etensk. .\kad. Handl. 1842: 17-,. Emendation. 
il'>-,5. Paradoxodoii Wagner, Schreijcr's Saugeth. Suppl. j: 803. Sorrx mrlanodon 

Blyth = Croctdura 1 Pachyura) nitidofulva Anderson. 
i8()7. Plerodus Schulze, Helios, Berlin, /.f 90. Croctdura suavolens Blasius (nee Pallas) 

— Sorex etrusciis Savi. 

4 species in the area covered by this list: 

Siiiicus dayi, page 69 
Sitncw, eiruseus, page 68 
Siiucin muruun. page 65 
Stincus it(dic.zkarius, page 69 
The retention ol' the genus Suncus, based on species which ha\c .m extra small 
a])|)<r unicuspid tooth, is largely a matter of convenience. Strictly speaking, it is not 


more than a subgenus oi' Crocidura. The Indian members of the genus were reviewed 
by Mrs. Lindsay, 1929, J. Bombay JV.H. Soc. 33: 326. This author recognizes an 
extraordinary number of species. There appear to be in the region now under discus- 
sion a pygmy species, for which the first name is etrusctts, a giant species, for which 
the first name is currently regarded as murinus, and a medium-sized group for which 
the first name is stoUczkanus. According to data from Lindsay, Miller and Bobrinskii, 
and examination of types and certain other specimens in London, the etruscus group 
contains forms which average as a rule 48 mm. or less in head and body length; the 
type of the Ceylon race and the form nitidofulvus are both a little larger (head and 
body 58 mm.). The Indian perrotteti and its allies has not to our knowledge been 
demonstrated to be other than racially separable from the European and South-^Vest 
Asian etruscus. The giant species, murinus, is very widely distributed in the tropics 
partly owing to human introduction, as it is a commensal species. Lindsay says the 
name murinus should be discarded as unidentifiable, and uses cneruleus for the giant 
house shrews, but murinus, which is used by Chasen and G. Alien, appears no more 
unidentifiable than several other very early names which are in current use for small 
mammals. From Lindsay's measurements, forms here referred to murinus average at 
least 93 mm. in head and body length, but the majority of specimens exceed 100 mm. 
The medium-sized group is much less common than the other two, and is confined 
to 'Western India. The head and body length in B.M. material averages 70-71 mm. 
Only one specimen examined for stoUczkanus is under 60 mm., and only one is over 
80 mm. The tail averages less than 70 per cent, of the head and body. Lindsay's 
measurements give an average of 70-73 mm. in head and body length for the group. 
S. dayi, which is little known, may well be a valid species. The type is darker than 
other specimens of the stoUczkanus group examined. The tail seems considerably less 
reduced, but unfortunately the type specimen does not bear measurements. In the 
type, the extra upper unicuspid, characteristic of the genus, is unusually large. In all 
probability it is a member of the stoUczkanus group. 

Suncus murinus Linnaeus, 1766 . House Shrew 

Approximate distribution of species: Philippines, Celebes, Borneo, Sumatra, Java, 
Bali, Malay States, to Annam, South-Eastern China, Formosa, Japan, Burma, west- 
wards to Kashmir, southwards to Ceylon; Arabia, Palestine, Egypt, Abyssinia, etc. 
Details of distribution apparently modified by human agency. 

Suncus murinus murinus Linnaeus, 1766 

1766. Sorex murinus Linnaeus, Syst. Nat. 12th ed. /.• 74. Java. 

1785. Sorex myosurus Pallas, Acta Acad. Sci. Petrop. 1781, 2: 337. Substitute for 
murinus Linnaeus. 

1792. Sorex caerulaeus Kerr, Anim. Kingd. 207. (Evidently a lapsus for caeruleus.) Java. 
(For status, see Chasen, 1940, Handlist Malaysian Mamm. 19.) 

181 1. Sorex indicus Gecjffroy, Ann. Mus. H.N. Paris, ij: 183. Pondicherry, India. 

1827. Sorex sonneralii Gcoffroy, Mem. Mus. H.N. Paris, 75.- 132. India. 

1 83 1. Sorex serpentarius Gcoflroy in Belanger, Voy. Indes Orient. Zool. 119. Pondi- 
cherry, India. 

1B45. Sorex nemorivagus Hodgson, Ann. Mag. N.H. ij: 269. Central region of Nepal. 




1859. Son-x iwin/wei Blyth, J. Asiat. Soc. Bengal, 1:8: 285. Amoy, Southern China. 

i860. Sorex albinus Blyth, J. Asiat. Soc. Bengal, 2g: 90, (nom. nud.). 

1870. Crocidura microhs Peters, Mber. Preuss. Akad. Wiss. 589. Hong Kong, China. 

1870. Crocidura [Pachyura) waldemarii Peters, loc. cit. 590. Bengal. 

1870. Crocidura (Pachyura) media Peters, loc. cit. 592. Paradcnia, Ceylon. 

(?) 1877. Crocidura (Pachyura) pealana Anderson, J. Asiat. Soc. Bengal, 46: 267. 
Sibsagar, Assam. 

1877. Crocidura (Pachyura) rubicunda Anderson, loc. cit. 277. Parcsnath Hill, cast of 
Hazaribagh^ Bihar, India. (Status j^(/t' Lindsay, 1929, 340.) 

1879. Crocidura andenoui Troucssart, Rev. Zool. Paris, 253. Khasi Hills, Assam. 

(?) 1881. Sorcx heddomd Anderson, Cat. Mamm. Ind. Mus. 179. Kollegal Hills, 
Coimbatore district, Southern India. 

(?) 191',. Crocidura imnchata Hatori, Taiwan Igakukai Zasshi, Ja]i. Number. (N.V.) 

Range: Lindsay quoted cacriileus from Gwalior, Central Provinces, Ximar, Bihar and 
Orissa. India, and Ceylon; according to Chascn, Lindsay's caeruleui = murinus, 
which he quotes from Malay Peninsula, Anambas, Sumatra, Java, Bali, Borneo. 
G. Allen quotes it from the larger towns of Southern C:hina, coastwise as far north 
as Fukien; Formosa, Hainan. Osgood recorded Suncus caerulcus from Annam, Indo- 
Clhina. Kuroda quotes the form ncinhoei from Formosa. 


1706. Sorcx piloridrs Shaw, Mus. Lever, i\- 31. .\(jt of Pallas, 1779, which is 

1800. Sorcx caeriilcsccin Shaw, Gen. Zool. Mamm. /.• 533. India (? Bengal). 
1831. Sorcx ,!;i,!iantcus Geoffroy, Voy. Belanger Indes Orient. Zcjol. 117. Bengal. 
Range: Darbhanga district, east of Nepal, and Midnapore district, India. 

Suncus murinus sacer Ehrenbcrg, 1833 

1833. Suncus sacer Ehrenbcrg, in Hemprich & Ehrenbcrg, Svmb. Ph\s. Manun. 2: 

folio k. Suez, Egypt. 

1834. Sorex crassicaudus Hemprich & Ehrenbcrg, in Liditciistcin's Darstellung 

Saugcth, pi. 40, tig. I, and text thereto. Neighbourhood of Suez, Egypt. 
1 808. Pachyura diiirrnnyt Fitzinger, S.B. Akad. Wiss. Wien. 1:56. Egypt. 
(?) 1935. Suncus tri\lrami B(jdenhcimer, Animal File in Pal<-stine, 9-,. Palestine. 

Su.NCUs .murinus socc.mus Hodgson, 1845 

1845. Sorcx soccatus Hodgson, Ann. Mag. N.H. ij: 270. Central region of Nepal. 
1855. Sorex heterodon Blyth, j. A^iat. Soc. Bengal, 24: 31. Clhcrrapunji, in Khasi HiUs, 


1850. Sorex montamis Kclaart, J. Ceylon Br. Soc. 2: 211. Pidurutal.igala, .\It. 

Nuw.ira I'.liya, Ceylon. 
1852. Sorex finiii^iueus Kcla.irt, J. Cleylon Br. .'\siat. Soc. _■.• 212. Dinibula, Nuwara 

Eliya, Ceylon. 
(?) 1855. Sorex kelaarli lUvth. J. .'\siat. Soc. Bengal, .■./; :;2. (uillc, C:c\lon. Based on 

a young spc( iincn ol iimnlanus according to I'hiilips ( 1935, Mamm. C.eylon). 



SuNCUS MURINUS GRiFFiTHi Horsfield, 1 85 1 

1 85 1. Sorex griffithi Horsfield, Cat. Mamm. Mus. E. India Co. 134. The label of the 
type has "Afghanistan", but this has been crossed out and "Silket" substi- 
tuted. See Lindsay (1929) on the suggestion that the type came from Assam. 

1877. Crocidura (Pachyura) blythii AnAtnon, ]. Asiat. Soc. Bengal, ^6, 2: 264. Assam. 

SuNCUS MURINUS NIGER Horsfield, 1 85 1 

1 85 1. Sorex niger Horsfield, Gat. Mamm. Mus. E. India Co. 135. Madras, India. 

(Elliot in MS.) 

SuNcus MURINUS KANDiANUs Kelaart, 1852 

1852. Sorex kandianus Kelaart, Prodr. Faun. Zeyl. 30. Ceylon. 

1870. Crocidura {Pachyura) ceylanica Peters, Mber. Preuss. Akad. \Viss. 591. Paradenia, 


1855. Sorex saturatior Hodgson, Ann. Mag. N.H. i6: i lo. Gangtok, Sikkim. 


1859. Sorex viridescens Blyth, J. Asiat. Soc. Bengal, 28: 285. Southern Malabar, 
India. Range: Madura and Trivandrum, Southern India. 


1859. Sorex lytleri 'Qlyth,]. Aiiiit. Soc. Bengal, 28: 285. Dehra Dun, Northern India. 
Range: Kumaon, Punjab, Kashmir. 


1877. Crocidura {Pachyura) fulvocinerea Anderson, J. Asiat. Soc. Bengal, ^G: 263. 
Gauhati, Assam. Range: North Kamrup and Valley of Assam. 

SuNcus MURINUS siNDENSis Andcrsou, 1877 

1877. Crocidura {Pachyura) sindetuis Anderson, J. Asiat. Soc. Bengal, ^6: 266. Karachi, 
Sind, India. Range includes Kathiawar, Rajputana and Cutch, India. 


1877. Crocidura {Pachyura) blanfordii Anderson, J. Asiat. Soc. Bengal, 46: 269. 
Khandalla, Western Ghats, 2,000 ft., India. Range includes Deccan area. 

SuNCus MURINUS RiuKiuANA Kuroda, 1924 

1924. Pachyura caerulea riukiuana Kuroda, On New Mamm. from Riukiu Islands, 

Tokyo, 3. Kinmu Kunchan, Okinawa, 200 ft., Liukiu Islands. Introduced 

Kiushiu, Japan. 

SuNcus MURINUS ZEYLANicus Phillips, 1928 

1928. Suncus zeylanicus Phillips, Spolia Zeylan, i^: 313. Gonagamma Estate. Kitul- 
gala, 900 ft., Ceylon. 



I9_'(). Suiicus nii^ir malabaricus Lindsay, J. Bombay N.H. Soc. 55; 334. No. -2437 from 
\'irajpct, .South Coore;, India, the only specimen to be mentioned by 
number, is assumed to be the holotype. Ranj^e: Coora; and Cochin, Southern 

Suncus etruscus Savi, 1822 Savi's Pygmy Shrew 

Apprdximate distribution of species: Southern Europe (Italy, Sicily, Sardinia, 
France, Spain, Greece, Hungary, etc.). Caucasus and Southern Russian Turkestan; 
Persia and P.ilestine iB.M.); recorded also from Algeria and Northern Nigeria (and 
quoted bv Boisrinskii from Arabia and Iraq;. As here understood, also Ceylon, India, north to Punjab, and Orissa, .Sikkim, Assam and Tenasscrim. A 
closelv .lUicd f)rm (or representative) occurs in the Malay States, and perhaps the 
species is represented also in East and South Africa. 


1822. Sorex etruscus Savi, Nuovo Giorn. de Lettcrati, Pisa, /; 60. Pisa, It.ily. 

183J. Sorcx pachvurui Kiister, Isis (Oken), 77. Cagliari, Sardini.i. 

i8j7. Crocidura siiaveolens Blasius, Saugeth. Deutschlands, 147. Not of Pallas, 181 1. 

Range: European range of the species; Persia, Turkestan, Ptdestine; Algeria and 
Nigeria ^sce Morrison-Scott, 1948, Mammalia, 10: 145). 


1842. Sorcx jhirultcli Du\ern(iy, Mag. Zool. Paris, 29. Nilgiri Hills, Southern 

(?) 1855. Sorcx hudgsoni Blyth, J. Asiat. Soc., i'_/.- 34. Darjeeling. 
1877. Crocidura (Pachvura) nilagirica Anderson, J. Asiat. Soc. Bengal, ^fi, 2: 274. 

Ootacamund, Nilgiri Hills, S(juthern India. 
1877. C.'rociduii! I'achvura) travancorcnsis .Anderson, loc. cit. 275. Tra\ancore, India. 

Range: Nilgiri Hills, Coori;, BcUary, etc., in Scjuthrrn India. 


1855. .So;<-v inicro/ivx Blyth, J. Asiat. Soc. Bengal, j?./.- 33. Landour, in Dehra Dun 
district, L'nited ProN'inces, Northern India. Range: Kumaon and probably 
Kangra, I' 

SuNCUS ETRUSCi:S MDIPES P)lyth, l8-,5 

18-,-,. .V()/,A iiudi/ici Blyth, J. .\siat. Soc. Bengal, _•./.■ 34. .Amherst, Tenasserim. 
?, 18-,-,. Siiiix iitralus Blyth, J. .Asiat. Soc. Bengal, i'^.- 34. Cherr.ipunji, Khasi Hills, 

?) 187",. I'acliviira a^\iitncii\i\ Anderson, I'.Z.S. 2';4. CJo.iljj.iroli, on Br.dimaputra. 
;?i 1877. (Jocidurn I'achxuKi] macruli\ .Anderson, |. Soc. Benf,'al, ^0, 2: 271. 

Ran^e includes |aiiiti.L Hills, .Assam and Shan States, Buinui. 



SuNCUS ETRUscus NiTiDOFULVus Andefson, 1877 

1877. Crocidura [Pachvura] mtidofulva Anderson, J. Asiat. Soc. Bengal, 46: 272. Lower 

Bengal, India. 
1855. Sorex nu'lanodon Blyth, J. Asiat. Soc. Bengal, 24: 33. Not of \Vagler, 1832. 
Range: Chaibassa, Orissa, India. 


1877. Crocidura {Pachyura) pygtnaeoides Anderson, J. Asiat. Soc. Bengal, 46: 279. 

1845. Sorex prgmaeiis Hodgson, Ann. Mag. N.H. i§: 269. Not of Laxmann, 1769. 
1867. Sorex hodgsoni ^erdon, Mamm. 57. Not of Blyth, 1855, which is a synonym of 

perrolteti according to Lindsay, 1929. 
Range: Darjeeling district, North-Eastern India. 


1932. Suncus fellowes-gordoni Philhps, Spolia Zeylan, ly: 124. \Vest Haputale, Ohiya, 
Central Province, Ceylon. 

Suncus stoliczkanus Anderson, 1877 Anderson's Shrew 

Appro-ximate distribution of species: India — Bombay, Central Provinces, Gwalior 
district, Rajputana, Kathiawar, Sind and Punjab. 

Suncus stoliczkanus stoliczkanus Anderson, 1877 

1877. Crocidura (Pachyura) stoliczkana Anderson, J. Asiat. Soc. Bengal, 46: 270. 

Bombay, India. 
1877. Crocidura [Pachyura) bidiana Anderson, loc. cit. 276. Madras, India. 
Range includes Gwalior, Salsette Island, Nimar and Hoshangabad, India. 

Suncus stoliczkanus subfulvus Anderson, 1877 

1877. Crocidura (Pachyura) subfulra Anderson, J. Asiat. Soc. Bengal, 46: 278. Cutch, 
India. Range includes Kathiawar and Sind. 

Suncus stoliczkanus leucogenys Dobson, 1888 

1888. Crocidura leucogenys Dobson, Ann. Mag. N.H. /.• 428. Ajmere (Rajputana 
district), India. 

Suncus dayi Dobson, 1888 Day's Shrew 

Approximate distribution of species: Southern Peninsular India. 

Suncus dayi Dobson, 1888 

1888. Crocidura dayi Dobson, Ann. Mag. N.H. /.• 428. Trichur, Cochin, India. (See 

Blanford, 1891, Fauna Brit. India, Mamm. 602.) Range includes Palni 

Hills, Southern India. 


i';tr; and Indian mammals 1738-1946 

Genus CROCIDURA \Vaglcr, 1832 

i8;j. Crocidura Waglcr, Isis, 275. (March, 1832.) Sonx leucodon Hermann. 

i8t)q. Leucodon Fatio, Faune Vert. Suisse, /.• 132. Sulsstitute for Crocidura. 

i8()7. Paurodus Schulze, Helios, Berlin, 14: 90. Sorcx arancus Schreber (not of Lin- 
naeus) = Sorcx russulus Hermann, and Sorex leucodon Hermann . 

1910. Hcliosorex Heller, Smith's Misc. Coll. jff, 15: (3. Heliosorex rooseieUi Heller, from 
East .\frica^ 

Of all genera in the class Mammalia, Crocidura must have been the largest collector 
of specific names. G. Allen has listed about no supposed species from Africa alone; 
we had on our preliminary lists 44 forms described binominally from the Palaearctic 
and Indian region; Chascn lists 29 more from the Malaysian region, and there are at 
least another 15 named from Celebes, the Philippines, Timor, etc. 

We have come to the conclusion that there are about 14 valid species in the region 
at present under discussion. It may be noted that Miller (1912) recognized three 
species occurring together in much of Western Europe [russula, leucodon and mimula = 
iuaveolens), and a fourth group of species, for which the prior name is caudata, from the 
.Mediterranean islands. Bobrinskii (1944) recognizes four species from the U.S.S.R. 
leucodon, russula, suaveolens and lasiura). G. Allen (1938) retained five species in China. 
His ilcnsis is the same as suaveolens; his two large species attenuata and dracula seem 
valid, although it is possible that dracula is not the prior name for the second large 
species, and his other two forms seem to be outlying races of russula. 

14 species in the area covered by this list: 

Crocidura attenuata, page 83 
Crocidura caudata, page 82 
Crocidura dracula, page 84 
Crocidura JJoweri, page 75 
Crocidura hispida, page 75 
Crocidura Iwrsfeldi, page 75 
Crocidura lasiura, page 84 
Crocidura leucodon, page 82 
Crocidura miya, page 75 
Crocidura olivieri, page 85 
Crocidura pergrisea, page 83 
Crocidura religiosa, page 75 
Crocidura russula, page 78 
Crocidura suaveolens, page 76 

Three species in the abo\e list of names have the tail longer than the head and 
bi)dv. This is a rare character in this genus. C. his/nda, i'rom the Andaman Islands, is 
a very large species (skull length about 27.7 mm.) kn(jwn by one specimen, which has 
the tail about 120 per cent, of the head and body. The elongated bristles on the tail 
which are characteristic of this genus and oi' Surum, but which are not invariably 
present in Crocidura, arc well developed. C. miya is a smaller species from Ceylon 
(skull length rciuglily 20 mm.!, with the t.iil aliout i i i per vcul. of the head and body. 


The caudal bristles are very reduced, but a few are traceable. C.floweri, from Egypt, 
is the third long-tailed species in the present region. The tail bristles seem absent in 
the specimens examined. Mr. R. \V. Hayman has remeasured the series on which the 
species was based, all of which are in spirit, and has obtained the following results: 

Head and 






















(Type) 54 




The condylobasal length varies between 17.8 and 19.2 mm., and the tail averages 
log per cent, of the head and body. 

All other species here dealt with have the tail shorter than the head and body. The 
only specimens examined in which it approaches this length are six skins labelled C 
attcnuata, from Upper Burma, which give an average of 98 per cent. 

There are three very small short-tailed species in the present region, in which the 
condylobasal length of the skull is not known to reach 18 mm. C. religiosa is an 
extremely small species from Egypt, in which the hindfoot is normally less than 
10 mm., the head and body length 45-55 mm., the tail relatively long (over 70 per 
cent, of the head and body), and the condylobasal about 15. 9-16. i mm. (B.M. speci- 
mens). In the Indomalayan region is a species which differs from religiosa by slightly 
larger average size (hindfoot not below 10 mm., head and body most often more than 
55 mm.). The caudal bristles in the specimens examined are traceable, though weak. 
The tail is long, more than 70 per cent, of the head and body. Shrews of this type 
have been examined from Ceylon ihorsfieldi), Indo-China {indochinensis) and Liukiu 
Islands [watasei). No characters which will distinguish these three forms specifically 
have been discovered, and horsfieldi is the prior name. According to its description, 
the form tadae appears to belong here. In this form the head and body can be as 
small as 50 mm. but the foot is at least iii mm., thereby differing from religiosa. 
Sixty-one millimetres is the highest measurement which has been noted for head and 
body length of horsfieldi and allies, and the tail can exceed 80 per cent, of the head 
and body. 

The widely distributed northern species, C. suaveolens, differs from horsfieldi and 
religiosa by its shorter tail, which is normally under 70 per cent, of head and body. 
The body length is approximately 55-75 mm., the tail percentage 43-63 per cent, in 
Europe, up to 67 per cent, in China. The only species from the British Islands 
{cassiteridum from the Scilly Isles) belongs here, and it is probable from descriptions 
that lignicolor may be a race (its skull is not fully known). C. suaveolens is represented 
in Palestine by portali and in North-West Africa by whitakeri. Two forms named 
recently by Goodwin from Persia may also be representati\'es of this species. In the 
remainder of the species, long series show condylobasal lengths of not under 18 mm. 
(with one individual exception). C. olivieri from Egypt stands apart from all the other 
short-tailed species in its unusually large size (condylobasal length 26.9-28.5 mm., 
B.M. material). This species looks like Simcus murinus, but has the dentition 



characteristic of Crociduia. It has shurt fur, the body length is 93-110 mm., the tail 
iwhich averages below 70 per cent, of head and body) is 63-70 mm. The Crocidura 
olivieii group (giant species) is common virtually throughout Africa, but absent in 
.\sia. The remainder have the condylobasal length normally between 18 and 
■2'-, mm. 

There seem to be about seven species definable in the russida group, medium-sized 
species with the tail shorter than the head and body, but the differences are average 
rather than absolute. C. russtila and C. leiicodon arc the earliest-named forms of this 
genus, both date from 1 780, and lussiila has line priority. These two species occur 
together, and dificr from each other in some colour details (see Miller, 191 2). In 
these species as here defined, the condylobasal length of the skull rarely reaches 
20 mm. For instance, in Miller's measurements, only two specimens oi leucodon out of 
33 noted are as much as 20 mm., and in nnsula 12 out of about 79 specimens reach 
20 mm. In the Turkish C. r. monacha, one in six reaches 20 mm. (B.M.). Two little- 
known forms which were named as races of riivsi/la, C. r. easpica from Persia and 
C. r. judaica from Palestine have the condylobasal length 2 1 mm. in the type skulls, 
and very likely represent C. lasiura, but before transferring them to that species more 
specimens will be needed. Bobrinskii has transferred the form C. leiicodon lasia to 
C. lasiura as a subspecies, and this seems necessary, as 12 duplicates for lasia have the 
condvlobasal length 20-23 mm., which is the normal size for lasiura. Miller's measure- 
ments for C. leucodon have the head and body 63-87 mm., the tail averaging about 
30-54 per cent, of it; and for russula head and body 64-95 mm., tail averaging 
,3_[j8 per cent, of it. There are other races in which the tail averages over 60 per cent, 
of head and body. These include C. russula cypria from Cyprus and C. r. caneae from 
Crete; also C. dsine-umi and allied forms from Japan, to which the forms vorax and 
rapax, described by G. Allen, from Yunnan, bear a close resemblance. As no 
characters have been found to separate the Mediterranean island forms cypria and 
caneae from the Japanese dsinezumi, the conclusion has been reached that it is wiser to 
call all these forms further outlying races oSI russula. The condylobasal length of 10 
specimens oi: dsinerumi in thi- H.M. varies between 18 and 19.4 mm., about the same 
size as published measurements fir caneae and cypria. Outlying forms, whii h 
apparently represent C. russula, are puUala from Kashmir and possibly sodyi from 


None of the firms just listed ha\e the tail as much as 70 per cent, ol'the head and 
bodv, which is characteristic of two species here retained, C. caudata [Mediterranean 
islands), and C. pernrisea (Kashmir and Baluchistan) (we have not seen fiergrisea, but 
from descriptions it is very like the Baluchistan firm, -arudnyi, which it antedates). 
These species have the condyfibasal length of the skull approximately 18-19.4 "ini. 
18-18.8 mm. in forms represented in London). The tail averages about 70-82 per 
cent., usually more than 70 per cent, of the head and body. A few specimens repre- 
senting caudata subspecies and zaruduyi in the B.M. indicate that the two species can 
be maintained on colour: zarudnyi is conspicuously paler both below and above; and 
pergrisea was described as very pale grey, below creamy white. The remaining forms 
in Asia are rather larger than russula, leucodon, caudata and pergrisca, although the 
difrercncc is an average one, the condylobasal length of the skull averaging at least 



20 mm. in each of the races. C. lasiura, from Manchuria, Ussuri, Asia Minor and the 
Caucasus, is a short-tailed species, with the tail fairly well haired; the condylobasal 
length in 24 specimens [lasiura, lasia) varies between 20 mm. and 22.6 mm., and the 
tail is relatively short, roughly 42-51 per cent, of the head and body length. In the 
form vamashinai (not represented in London, but here tentatively regarded as a race), 
the skull length is 23.5-25 mm., but smaller specimens seem covered by larger speci- 
mens oi lasiura. (Kuroda gives measurements of 21-24 mm. for lasiura^) The body 
length is 73-98 mm. in more typical forms, but can be as much as 1 12 mm. in larger 
specimens of vamashinai. Two species, which are mostly Indomalayan in distribution, 
have the large skull o{ lasiura but differ in having the tail at least 60 per cent., usually 
over 70 per cent, of head and body. (Allen's measurements for attenuata have the tail 
averaging about 64 per cent., but most of our specimens are over 70 per cent.) Two 
named races of dracula have the tail 60 per cent, in the types, but it is more usual for 
the tail to exceed or approximate 80 per cent, in this species. The two species occur 
together, and the prior names seem to be attenuata and dracula. The Himalayan forms, 
rubricosa and kingiana, seem to represent attenuata. Twelve skulls of attenuata (B.M.) 
have the condylobasal length ig. 8-22.1 mm., but only once under 20 mm. All G. 
Allen's specimens exceed 20 mm. Twelve specimens in the B.M. representing 
rubricosa and kingiana have the condylobasal length 19.3-23.9 mm., but only once 
over 22 mm. and twice under 20 mm. From descriptions, the Formosan form tanakae 
should represent attenuata. 

The second large species in Southern China and Northern Burma is C. dracula, 
unless this represents one of the numerous earlier-named forms from the Malaysian 
region. Where it occurs with attenuata it is larger on average. The type of the race 
grisescens has the greatest length of skull only 2 1 .6 mm. and possibly does not represent 
the species; otherwise no specimen with the skull length less than 22 mm. has been 
noted. The body length is about 84-105 mm., and the largest skulls are about 
24.3 mm. in length. All species dealt with above have the caudal bristle hairs at least 
perceptible, except apparently floweri. Of the specimens seen, they were noted as 
being most reduced in rubricosa, horsfieldi and miya. 

These results, which must be regarded as provisional, can be arranged in kev form, 
as follows; 

1. Tail clearly exceeds length of head and body. 2 

Tail shorter than head and body. 4 

2. Length of skull 27.7 mm. Tail more than 120 per cent, of head and body. 


Length of skull about 20.5 mm, and less. Tail 109-1 1 1 per cent, of head and 

body. 3 

3. Bristles on tail barely perceptible. Length of skull about 17. 8-19. 2 mm. Head 

and body 54 mm.,' and less. (Type in B.\L) CROCIDURA FLO]\'ERI 

Bristles on tail perceptible. Skull length approximately 20.5 mm. Head and bodv 
79 mm. (type specimen, B.M.) CROCIDURA MIl'A ' 


pai.aearc;tic and Indian mammals 1758-1946 

4. Small; condylobasal length of skull less than 18 mm. 5 

Larger; condylobasal length of skull normally at least i8 nnn. 7 

5. Tail shorter, averaging less than 70 per cent, of head and body. 

Forms examined: whilaken, mtmiila, ilensis, coreac -- s/iariluiigensis, portali, 
Tail longer, averaging over 70 per cent, of head and body. 6 

6. Hindl'oot normally 8?,-9 mm. (Egypt). 

CROCIDL'RA RELIGIOSA (Several specimens a\ailable for examination) 
Hindfoot 10 mm., and more (Indomalayan). CROCIDURA HORSEIEI.DI 
Forms examined: hnisficldi, indochinensis, ivatasei. 

-. Verv large species, condylobasal length 26.9-28.5 mm. (Fur short; appearance 
reminiscent of a house-dwelling form.) 

CROCIDURA OLIVIERI (Several specimens available for examination) 
Smaller species; length of skull not known to exceed 25 mm. 8 

8. In the majoritv of specimens, the skull is less than 20 mm. in length. 9 

In the majority of specimens, the skull is at least 20 mm. in length. 12 

C). Tail long, averages at least 70 per cent, of head and body length. 10 

Tail shorter, a\erages less than 70 per cent, of head and body length. 1 1 

10. Colour paler abo\e and below. 

CROCIDURA PERGRISEA 'Kashmir, Baluchistan) 
Form examined : znrudnyi. 
Colour darker above and below. 

CROCIDURA CAUDATA (Western Mediterranean) 
Forms examined: halearica, cyrnensis. 

11. Di\ision of colour between light underparts and flark back gencrallv more 


Forms examined : leiicodoii, siciila, pcrsica. 
Division of colour between underparts and back usually less abrupt. (The 
differences between these two species, which occur together, arc not very 
strongly marked.) CROCIDURA RUSSULA 

Forms examined: nisuilii, duneziimi, iimbnna, crpria^ monacha, c/iisai, caspica, 
ciiilrae, piilchra, cancae, judaica, pi'ln, \ndyi. The forms jiidaica and ca\pica seem 
based iin one specimen each with a skull too lart;e for riisMihi, and further 
material m.iy slmw them to be representatives of C lasiiira. 

12. Tail relati\elv short, approximatinij; half head and bodv length, nv less. 

Forms examined: lasiiira, la.\ia, thoman. 
Tail long, rarely as low as 60 per cent., iiKJstly exceeding 70 per cent, of head 
and body. 13 



13. In the majority of specimens the length of the skull is less than 22 mm. 

Forms examined : attenuata, ruhricosa, kingiana. 
In the majority of specimens the length of the skull is 22 mm., and more. 

Forms examined: dracida, praedax = dracula. 

Crocidura hispida group. (Very long-tailed species.) 

Crocidura hispida Thomas, 191 3 Andaman Island Spiny Shrew 

Approximate distribution of species: Middle Andaman Island, Bay of Bengal. 

Crocidura hispida Thomas, 1913 

1913. Crocidura hispida Thomas, Ann. Mag. N.H. //.• 468. Northern end Middle 
Andaman Island, Andaman Islands. 

Crocidura floweri Dollman, 191 5 Flower's Shrew 

Approximate distribution of species: Egypt. 

Crocidura floweri Dollman, 19 15 

1915. Crocidura floweri Dollman, Ann. Mag. N.H. i^: 515. Gizeh, Egypt. See also 
1916, Ann. Mag. N.H. ly: 192. 

Crocidura miya Phillips, 1929 Ceylon Long-tailed Shrew 

Approximate distribution of species: Ceylon. 

Crocidura miya Phillips, 1929 

1929. Crocidura miya Phillips, Spolia Zeylan. i^: 113. Moolgama, village in the 
Nilambe district of Kandyan Hills, about 3,000 ft., Ceylon. 

Crocidura suaveolcns group. (Pygmy species.) 

Crocidura religiosa GeofTroy, 1827 Egyptian Pygmy Shrew 

Approximate distribution of species: Egypt. Probably also represented iit Somali- 
land, Uganda, Sudan, under the names nana, nanilla, pasha. 

Crocidura religiosa Geoffroy, 1827 

1827. Sorex religiosus I. Geoffroy, Mem. Mus. H.X. Paris, /j.- 128. Types mummified, 
Thebes, Egypt. 

Crocidura horsfieldi Tomes, 1856 Horsfield's Shrew 

Approximate distribution of species: Ceylon, also apparently represented in 
Kashmir, Indo-China, Siam (Tate), Northern Burma and Liukiu Islands. 


Crocidura horsfieldi horsfieldi Tomes, 1856 

1856. Sorex /wrsfieldii Tomes, Ann. Mag. N.H. ly: 23. Ceylon. 

1870. Crocidura retiisa Peters, Mber. Preuss. Akad. Wiss. 585. Paradenia, Ceylon. 

Crocidur.-\ (.') HORSFIELDI .MvoiDEs Blanford, 1875 

1875. Sorex [Crocidura) myoides Blanford, J. Asiat. Soc. Bengal, ./^, 2: 106. Leh, 
Ladak. From descriptions seems nearest to horsfieldi. 

Crocidura horsfieldi indochinensis Robinson & Kloss, 1922 

1922. Crocidura indochinensu Robinson & Kloss, /Xnn. Mag. N.H. g: 88. Dalat, Lang- 

bian Plateau, 5,000 ft., Annam, Indo-China. Range includes Northern 

Burma; and Siam, according to Tate. 

Crocidur,.\ horsfieldi watasei Kuroda, 1924 

1924. Crocidura ivatasei Kuroda, New Mamm. from Riukiu Islands, Tokyo, i. Komi, 
Amamioshima, Liukiu Islands. 

Crocidura horsfieldi tadae Tokuda & Kano, 1936 

1936. Crocidura tadae Tokuda & Kano, Annot. Zool. Jap. i§: 429. Koto-sho = 
Island of Botel Tobago (east of Formosa). 

Crocidura suaveolens Pallas, 181 1 Lesser White-toothed Shrew 

Appniximatc distribution of species: Germany, southwards to Yugoslavia, Bul- 
garia and Greece; France, Switzerland, south to Italy and Sardinia; Spain; 
represented in the Scilly Islands; Central and Southern Russia (north to southern 
districts of Moscow Province), Estonia; Russian Turkestan, and Ussuri district of 
Eastern Siberia; Sinkiang, Mongolia, Korea, most of the larger states of China from 
Shantung and Chckiang, westwards to Szechuan; Persia, Palestine; Morocco and 
Algeria. Probably also ranges in East and South-West Africa under the name hicolor. 

Crocidur.'\ suaveolens suaveolens Pallas, 181 1 

181 1. Sorex suaveolens Pallas, Zoogr. Ross. As. /.• 133, pi. 9, fig. 2. Khersones, Crimea, 

Southern Russia. 
(.'') 1934. Crocidura suaveolens mordeni Goodwin, Amer. Mus. Nov. No. 742: i. Tuz 

Rulak, 50 miles north of Kizil Orda (Perovsk), Kazakstan, 600 ft., Russian 

C'entral Asia. 
Range: Lowlands of Russia and Russian Central Asia. 

Crocidura suaveolens whitakeri de \Vinton, 1898 

1898. Crociduia whitakeri de Winton, P.Z.S. i8gy: 954. Sierzet, half-way between 
Morocco City and Mogador, Morocco. 

Crocidura suaveolens lignicolor Miller, 1900 

1900. Crocidura lignicolor Miller, Proc. Washington Acad. Sci. 2: 39. Jungle east of 
Maralbashi, near Yarkand River, Chinese Turkestan. The skull of this form 
seems not fully known, but the external measurements suggest a small form 
of the present species. 



Crocidura suaveolens mimula Miller, 1901 

1 90 1. Crocidura mimula Miller, Proc. Biol. Soc. Washington, 14: 95. June, 1901. 
Ziiberwangen, St. Gallen, Switzerland. 

(?) 1839. Crocidura aranea var. minor de Selys Longchamps, Etudes de Micromamm. 
35. Silesia. 

1901. Crocidura antipae Matschie, S.B. Ges. Naturf. Fr. Berlin, 228. November, 1901. 
Siulnita and Barza, Rumania. 

igo2. Crocidura minuta Lydekker, Zool. Record, igoi, Mamm. : 27. Accidental re- 
naming of mimula. 

1 92 1. Crocidura dinnicki Ognev, Ann. Mus. Zool. Acad. St. Petersb. 340. Stavropol. 
Northern Caucasus. (Status ^rf^ Bobrinskii.) 

Range : France, Germany, Bohemia, Transylvania, Yugoslavia, Rumania, Bulgaria, 
Switzerland, Italy, Greece, Caucasus. 

Crocidura suaveolens ilensis Miller, 1901 

1901. Crocidura ilensis Miller, Proc. Biol. Soc. Washington, 14: 157. Kukturuk, Hi, 

5,400 ft., extreme W'estern Chinese Turkestan. Ranges to Mongolia. 

Bobrinskii thinks this is probably a synonym oi suaveolens. 

Crocidura suaveolens shantungensis Miller, 1901 

1 90 1. Crocidura shantungensis Miller, Proc. Biol. Soc. Washington, i.}: 158. Chimeh, 
Shantung, China. 

1907. Crocidura coreae Thomas, P.Z.S. igo6: 860. Mingyong, no miles south-east of 

Seoul, Korea. 
1927. Crocidura longicauda Mori, J. Chosen N.H. Soc. 5.' 28. Seoul, Korea. 
Range: Korea, Shansi, Shensi, Shantung, Chekiang, in China; Tsushima I. 

Crocidura suaveolens iculisma Mottaz, 1908 

1908. Crocidura mimula iculisma Mottaz, Bull. Soc. Zool. Geneve, /.• 119. Lignieres- 

Sonneville, Charente, France. 

Crocidura suaveolens cantabra Cabrera, 1908 

1908. Crocidura cantabra Cabrera, Bol. Soc. Esp. H.N. 8: 239. Basque Provinces, 
Spain (exact locality unknown). 

Crocidura suaveolens italics Cavazza, 191 2 

1912. Crocidura mimula italica Cavazza, Boll. Mus. Zool. Anat. Comp. Torino, 27, 
653: 12. Delia Valle Padana, Italy. 

Crocidura suaveolens sarda Cavazza, 191 2 

191 2. Crocidura sicula var. sarda Cavazza, Boll. Mus. Zool. Anat. Comp. Torino, 27, 
659: 7. Cagliari, Sardinia. 

Crocidura suaveolens portali Thomas, 1920 

1920. Crocidura portali Thomas, Ann. Mag. N.H. 5.- 1 19. Ramleh, south-east of Jaffa, 



Crocidura suaveolens orientis Ognev, 192 1 

1 92 1. Crocidura suaveolens orientis Ognev, Ann. Mus. Zool. Acad. St. Petersb. 22: 341. 
Ncbilmi, valley of River Tuman-gan, Ussuri region of Eastern Siberia. 

Crocidur.x suaveolens phaeopus G. Allen, 1923 

1923. Crocidura ilcnsis phaeopus G. Allen, Amer. Mus. Nov. No. 100, 7. Wanhsien, 

Szcchuan, C^hina. Range includes Hupch and Southern Shensi, China. 

Crocidura suaveolens cassiteridum Hinton, 1924 

1924. Crocidura cassiteridum Hinton, Ann. Mag. N.H. 14: 509. An uninhabited island, 

Scilly Islands '(off Cornwall, England). 

Crocidura suaveolens debeauxi Dal Piaz, 1925 

1925. Crocidura mimula debeauxi Dal Piaz, Atti Soc. Vcn. -Trent. Sci. Nat. 16 (sep. 

pag). Frugarolo, Prov. de Allessandria, Northern Italy. 

Crocidura suaveolens lar G. Allen, 1928 

1928. Crocidura lar G. Allen, Amer. Mus. Nov. No. 317: i. Tsagan Nor, Central 
Gobi, Mongolia. 

Crocidura (?) suaveolens hyrcani.'^ Goodwin, 1940 

1940. Crocidura hyrcania Goodwin, Amer. Mus. Nov. No. 1082; i. Turkman plains, 

about 60 km. north-east of Astrabad, on banks of the Gurgan River, sea 

level, Persia. 

Crocidura (?) suaveolens astrabadensis Goodwin, 1940 

1940. Crocidura astrabadensis Goodwin, Amer. Mus. Nov. No. 1082: 3. Dar Kaleh, 
about 40 km. east of Astrabad, sea level, Persia. 

Crocidura (?) suaveolens oyaensis Heim dc Balsac, 1940 

1940. Crocidura oyaensis Heim de Balsac, C.R. Acad. Sci. Paris, 211: 296. Yeu Island, 
off Vendee, Western France. 

Crocidura russula group. (Medium-sized species.) 

Crocidura russula Hermann, 1780 Common European White-toothed Shrew 
Approximate distribution of species: France, Channel Islands, Sardinia, Corsica, 
Switzerland, Italy, Spain, Portugal, Belgium, Holland, Germany, Poland, Crete; 
Caucasus and Southern Russian Turkestan, east to Pamir Mountains; Asia Minor, 
Persia, Palestine, Afghanistan (B.M. specimens collected by Chaworth-Musters 
identified as this species); Kashmir; Japan; Yunnan, China; ? Korea; Morocco, 
Algeria, Tunis. Probably also represented in Kenya, Sudan, Angola, etc. 

Crocidura russula russula Hermann, 1 780 

1780. Sorex russulus Hermann, in Zimmermann, Geogr. Gesch. 2: 382. Near Stras- 
bourg, Bas-Rhin, Eastern France. 



1778. Sorex araneus Schreber, Saugeth. j; 573. Not of Linnaeus, 1758. 

(?) 1780. Sorex constrictus Hermann, in Zimmermann, Geogr. Gesch. 2: 383. Near 

Strasbourg, France (based on young animal). 
(?) 1792. Sorex unicolor Kerr, Anim. Kingd. 208. Strasbourg, France. 
1798. Sorex musaraneus Cuvier, Tabl. Elem. H.N. des Anim. 109. France. 
(?) 1800. Sorex leueums Shaw, Gen. Zool. /, 2: 538. Strasbourg, France. 
1801. Sorex araneus cinereus Bechstein, Gemeinn. Nat. Deutschlands, /, 2nd ed.: 867 

(misprinted 863). Thuringia, Germany. 
1 801. Sorex araneus eandidus Bechstein, loc. eit. Thuringia, Germany. 
1832. Sorex Jimbriatus Wagler, Isis, 54. Bavaria, Germany. 
1832. Crocidura moschata Wagler, Isis, 275. Substitute for Sorex Jimbriatus. 
1832. Crocidura major Wagler, Isis, 1218. Bavaria, Germany. 
1832. Crocidura rufa Wagler, Isis, 1218. Banks of Rhine, Germany. 
1832. Crocidura poliogastra Wagler, Isis, 1218. Banks of Rhine, Germany. 
1832. Sorex thoracicus Savi, Nuovo Giorn. de Letterati, Pisa, 24: 52. Near Pisa, Italy. 
(?) 1839. Sorex inodorus de Selys Longchamps, Etudes Micromamm. 34. Silesia. 
(?) 1839. Crocidura aranea var. albiventris de Selys Longchamps, loc. cit. No locality. 
(?) 1839. Sorex hydruntina Costa, Fauna del Ragno di Napoli, Mamm. 6. Otranto, 

Calabria, Italy. 
1855. Sorex chrysothorax Dehne, Allg. Deutsche Naturhist. Zeitung, /; 241. Wilsdurf, 

near Dresden, Germany. 

Range: Holland, Belgium, France, Germany, Switzerland, Italy. Bobrinskii thinks 
the next is the same and includes Russia, Caucasus and Northern Persia in the 

Crocidura russula GtJLDENSTAEDTi Pallas, 181 1 

181 1. Sorex guldenstaedtii Pallas, Zoogr. Ross. Asiat. /.• 132, pi. 9, fig. 1. Near Dushet, 

Georgia, Transcaucasia. 
(?) 1863. Sorex {Crocidura) fumigatus de Filippi, Arch. Zool. Anat. Fisiol. Genova, 

2: 379. Tehran, Northern Persia. Range includes Simla, according to 

1889. Crocidura longicaudata Tichomirov & Kortchagin, Mem. Soc. Amis. Sci. Nat. 

Moscou, §6, 4, 1 : 17. Sukhum, Black Sea, Russia. 
1889. Sorex bogdanowii Tichomirov & Kortchagin, loc. cit. 
1914. Crocidura russula aralychensis Sa.tVinm,'\i\\.t. Kaukas. Mus. 8: 92. Marshy shores 

of River Karasu, near Aralych, Caucasus. 

Crocidura russula dsinezumi Temminck, 1844 

1844. Sorex dsi-nezumi Temminck, in Siebold, Faun. Japon. Mamm. 26. Kiushiu, 

1844. Sorex kinezumi Temminck, loc. cit. 26 (footnote). 

1845. [Sorex) kinezumi Temminck, in Siebold, Fauna Japon. Mamm. ^, Tabl. iv, figs. 

6-1 ic. 
Range: Kiushiu, Shikoku, ? Oki Is. 

crocidura russula umbrina Temminck, 1844 

1844. Sorex umbrinus Temminck, in Siebold, Faun. Japon. Mamm. 27. Miyanoura, 
Yakushima, Japan (Kuroda). 



Crocidura russula agilis Levaillant, 1867 

1867. Pachvura agilis Levaillant, in Loche, Explor. Sci. de I'Algerie, Zool., pi. 4, 
fig. '2. Algeria. ("The atlas of this work was published many years before the 
text according to Cabrera." G. Allen (who dates the name from 1850).) 

(?) 1856. Sorex mmintanmis Pomel, C.R. Acad. Sci. Paris, ^2: 653. Algeria. [N.V.)^ 

1867. Pachnira pigmaea Loche, Explor. Sci. de I'Algerie, Zool. 88. Ain-el-Bel, Algeria. 

Range: Morocco to Tunis. 

Crocidura russula cypria Bate, 1904 

1904. Crocidura russula cypria Bate, P.Z.S. igo3, 2; 344. Cyprus. 

Crocidura russula monacha Thomas, 1906 

1906. Crocidura russula monacha Thomas, Ann. Mag. N.H. 77.- 417. Scalita, near 
Trebizond, 700-1,000 m., Asia Minor. 

Crocidur.'v russula chisai Thomas, 1906 

1906. Crocidura dsi-nezumi chisai Thomas, P.Z.S. igo^, 1 : 340. Tsunagi, near Morioka, 

Northern Hondo, Japan. 

Crocidura russula caspica Thomas, 1907 

1907. Crocidura russula caspica Thomas, Ann. Mag. N.H. 20: 197. Southern coast of 

Caspian Sea, Northern Persia. It is possible that this little-known form 
represents C. lasiura. 

Crocidura russula pulchra Cabrera, 1907 

1907. Crocidura russula pulchra Cabrera, Ann. Mag. N.H. 20: 213. Valencia, Spain. 

Range; Portugal (part) ; Spain (Central and Southern) ; lowlands of France, 

south of the Gironde. 

Crocidura russula cintrae Miller, 1907 

1907. Crocidura russula cintrae Miller, Ann. Mag. N.H. 20: 390. Cintra, near Lisbon, 

Crocidura russula caneae Miller, 1909 

1909. Crocidura caneae Miller, Ann. Mag. N.H. 3: 418. Crete. 

Crocidura russula pullata Miller, 191 1 

191 1. Crocidura pullata Miller, Proc. Biol. Soc. ^Vashington, 24: 241. Kotihar, 

7,000 ft., Kashmir. 

Crocidura russula ichnusae Festa, 1912 

191 2. Crocidura ichnusae Festa, Boll. Mus. Zool. Anat. Comp. Torino, 27, 648: i. 

Piscina, Lanusei, .Sardinia. 

Crocidura russula mimuloides Cavazza, 1912 

191 2. Crocidura russula mimuloides Cavazza, Boll. Mus. Zool. Anat. Comp. Torino, 
-7' *^53' "^l- B"gg'o'o> Ticino Alps, Italy. 


Crocidura russula yebalensis Cabrera, 191 3 

1913. Crocidura yebalensis Cabrera, Bol. Soc. Esp. H.N. 13: 400. Tetuan, Morocco. 

Crocidura russula judaic a Thomas, 1919 

1919. Crocidura russula judaica Thomas, Ann. Mag. N.H. 2' 32. Near Jerusalem, 
Palestine. It is possible that this little-known form represents C. lasiura. 

Crocidura russula peta Montagu & Pickford, 1923 

1923. Crocidura russula peta Montagu & Pickford, P.Z.S. 1044. Guernsey, Channel 

Crocidura russula vorax G. Allen, 1923 

1923. Crocidura vorax G. Allen, Amer. Mus. Nov. No. 100: 8. Timber-line forest on 
Ssu Shan (Snow Mountain), Likiang Range, 12,000 ft., Yunnan, China. 

Crocidura russula rapax G. Allen, 1923 

1923. Crocidura rapax G. Allen, Amer. Mus. Nov. No. 100: 9. Yinpankai, Mekong 

River, 9,000 ft., Yunnan, China. 

Crocidura russula intermedia Kuroda, 1924 

1924. Crocidura dsi-nezumi intermedia Kuroda, New Mamm. from Riukiu Islands, 

Tokyo, 2. Nishino-omote, Tanegashima Island, 200-400 ft., Japan. 

Crocidura russula orii Kuroda, 1924 

1924. Crocidura dsi-nezumi orii Kuroda, New Mamm. from Riukiu Islands, Tokyo, 3. 
Komi, Amamioshima, Liukiu Islands. 

Crocidura russula pamirensis Ognev, 1928 

1928. Crocidura pamirensis Ognev, Mamm. E. Europe, N. Asia, /.• 366. Near Lake 

Drum, south slope Pamir Range, 12,000 ft., Russian Asia. 

1929. Crocidura serezkjensis Laptev, Mater. Centr. Asiat. Zool. Gard. /.• 16. [N.V.) 

Ognev, Mamm. E. Europe, N. Asia, 2: 771. Lake Severskoe, Pamir 

Crocidura russula ^uelpartis Kuroda, 1934 

1934. Crocidura dsi-nezumi quelpartis Kuroda, J. Mamm. /j.- 236. Seikiho, Quelpart 
Island, off Korea. 

Crocidura (?) russula sodyi Kuroda, 1935 

1934. Crocidura neglecta Kuroda, J. Mamm. 13: 238. Not of Jentink, 1888. 

1935. Crocidura sodyi Kuroda, Zool. Mag. Tokyo, 4y: 327. To replace neglecta 

Kuroda, preoccupied. Bampo, Korea. (Size large, skull of type, 20 mm. But 
possibly represents russula, as it occurs with the much larger C. lasiura 
yamaskinai which has the same type locality.) 

Crocidura (?) russula corsicana Heim de Balsac & Reynaud, 1940 
1940. Crocidura corsicana Heim de Balsac & Reynaud, Bull. Soc. Zool. France, 6^: 
216. Francardo, He Rousse, Corsica. 



Crocidura russula anthonyi Heim de Balsac, 1940 

1940. Crocidura anthonvi Hcim de Balsac, Bull. Mus. H.N. Paris, 12: 382. Gafsa, 

Croc:idur.\ (?) russula foucauldi Agacino, 1943 

1943. Crocidura foucauldi Agacino, Bol. Soc. Esp. H.N. 41: 37. Isaguen, 1,500 m., 
Beni Seddat, Rif, Spanish Morocco. 

Crocidura leucodon Hermann, 1780 Bicolor White-toothed .Shrew 

Approximate distribution of species: Holland, Belgium, France, Germany, Poland, 
Switzerland, Italy, apparently represented in Sicily; Yugoslavia, Transylvania; 
Central and Southern Russia, including Crimea, Caucasus, Eastern Turkestan, and 
northwards to Central Siberia (Minussinsk steppe, Semipalatinsk province); Persia. 

Crocidura leucodon leucodon Hermann, 1780 

1780. Sorex leucodon Hermann, in Zimmermann, Gcogr. Gesch. 2: 382. \'icinity of 

Strasbourg, Bas Rhin, Eastern France. 
1791:. Sorex albipes Kerr, Anim. Kingd. 208. 

i8Gq. Leucodon microurus Fatio, Faune Vert. Suisse, /.■ 137. Substitute for leucodon. 
1807. Crocidura leucodus Schulze, Helios, Berlin, /./.■ 90. Substitute for leucodon. 
Range: European and Russian range of the species, except Sicily. 

CiRociDUR.A '?) leucodon sicula Miller, 1901 

Kioi. Crocidura sicula Miller, Proc. Biol. Soc. Washington, 14: 41. Palermo, Sicily. 

[Crocidura sicula Giglioli, 1879, Aich. Xaturgesch. /.• 96, nom. mid.) Perhaps 

a form of C. russula. 

C.ROCiDURA leucodon persica Thomas, 1907 

1907. Crocidura leucodon persica Thomas, Ann. Mag. X.H. 20: 198. Elburz Mountains, 

near Dcmavend, 6,500 ft., Persia. 

1908. Crocidura leucodon caspica Lydekker, Zool. Record, igoy, Mamm.: 59. Acci- 

dental renaming of C. I. persica. 

Crocidura leitiodon narent.-\e Bolkay, 1925 

1925. Crocidura leucodon narentae Bolkay, X<>\it. Mus. Sarajevo, /.- 7. Between Capljna 
and the uld Roman defensive castle, Mcigorjelo, Herzegovina, \'ugoslavia. 

Crocidur.\ leucodon sibirica Dukelski, 1930 

1930. Crocidura leucodon sibirica Dukelski, Zool. Anz. 88: 75. Village of Osnatschen- 
noje, on River Yenesei, 96 km. south of Minussinsk, Siberia. Bobrinskii 
calls this form C. I. myoides fBlanfordj, but myoides Blanford from description 
seems to be a form of C. horsjieldii. The present name is available for the 
Siberian race. 

Crocidura caudata .Miller, K|0i .Mediterranean Long-tailed Shrew 

.Approximate di^triljutidii nf species: Sicily, Corsica, Balearic Islands. 



Crocidura caudata caudata Miller, igoi 

igoi. Crocidura caudata Miller, Proc. Biol. Soc. Washington, 14: 42. Palermo, Sicily. 

Crocidura caudata cyrnensis Miller, 1907 

1907. Crocidura cyrnensis Miller, Ann. Mag. N.H. 20: 390. Bastia, Corsica. 

Crocidura b.alearic.a Miller, 1907 

1907. Crocidura balearica Miller, Ann. Mag. N.H. 20: 391. San Cristobal, Minorca, 
Balearic Islands. 

Crocidura pergrisea Miller, 191 3 Pale Grey Shrew 

Approximate distribution of species: Kashmir, Baluchistan and Eastern Persia. 

Crocidura pergrisea pergrisea Miller, 191 3 

1913. Crocidura pergrisea Miller, Proc. Biol. Soc. Washington, 26: 1 13. Skoro Loomba, 
Shigar, Baltistan, 9,500 ft., Kashmir. 

Crocidura pergrisea zarudnyi Ognev, 1928 

1 92 1. Crocidura tatianae Ognev, Ann. Mus. Zool. Acad. St. Petersb. 22: 338. Eastern 

Persia (Baluchistan). Not C. tatiana Dollman, 1915. 
1928. Crocidura zarudnyi Ognev, Mamm. E. Europe, N. Asia, /.• 341. New name for 

tatianae Ognev nee Dollman. 
Specimens examined from Kelat and Turbat, Indian Baluchistan. 

Crocidura atienuata Milne-Edwards, 1872 -Grey Shrew 

Approximate distribution of species: China, states of Kiangsu, Chekiang, Hupeh, 
Szechuan, Hunan, Fukien, \Vestern Yunnan ; Hainan, Northern Burma, Assam, 
Bhutan Duars, Sikkim, Kumaon, Punjab, Kashmir; apparently Formosa. 

Crocidura attenuata attenuata Milne-Edwards, 1872 

1872. Crocidura attenuata Milne-Edwards, Rech. H.N. Mamm. 263, pi. 38B, fig. i, 

pi. 39A, fig. 2. Moupin, Szechuan, China. 
1926. Crocidura grisea Howell, Proc. Biol. Soc. \\'ashington, 29- '37- Seventy-five 

miles south-west of Yenpingfu, 500 ft., Fukien, China. 
Range: China, as listed above, and including Hainan; Northern Burma (B.M.) 

Crocidura attenuata rubricosa Anderson, 1877 

1877. Crocidura rubricosa Anderson, J. Asiat. Soc. Bengal, 46, 2 : 280. Sibsagar, Assam. 

Specimens examined from Assam, Kamrup, Bhutan Duars, and Pashok, 

near Darjeeling. 

Crocidura attenuata kingiana Anderson, 1877 

1877. Crocidura kingiana Anderson, J. Asiat. Soc. Bengal, 46, 2: 281. Sikkim. Speci- 
mens examined from. Kumaon and Punjab. 

Crocidura (?) attenuata tanakae Kuroda, 1938 

1938. Crocidura tanakae Kuroda, Handlist Jap. Mamm. 81. Shohosha, Horigai, 
Taichusiu, Central Formosa. 



Crocidura dracula Thomas, 1912 Dracula Shrew 

Approximate distribution of species: Yunnan, Fukien, Northern Burma, Indo- 
China. The form dracula requires comparison with C. baluensis Thomas, 1898, Borneo 
(? = a race of C. oricntalis ]cn\.\nk, 1890, Java), and other earher-named extralimital 

Crocidura dracula dracula Thomas, 191 2 

1912. Crocidura dracula Thomas, Ann. Mag. N.H. g: 686. Probably near Mongtze 

(Mengtsz), Southern Yunnan, China. 
1923. Crocidura praedax Thomas, Ann. Mag. N.H. //.■ 656. LitLiang Valley, Central 

Yunnan, 9,500 ft., China. 

Crocidura dracul.^ grisescens Howell, 1928 

1928. Crocidura grisescens Howell, J. Mamm. g: 60. Kuatun, Fokien, South-Eastern 

Crocidura dracul.'\ mansumensis Carter, 1942 

1942. Crocidura dracula mansumensis Carter, Amer. Mus. Nov. No. 1208: i. Mansum 
(25.47' N., 96.16° E.), 3,200 ft.. Northern Burma. 

Crocidura lasiura Dobson, 1890 Ussuri Large White-toothed Shrew 

Approximate distribution of species: Ussuri region of Eastern Siberia, Manchuria, 
Korea; Kiangsu, in China. Asia Minor and Caucasus, and Northern Persia according 
to Bobrinskii. 

Crocidura lasiura lasiura Dobson, 1890 

1890. Crocidura lasiura Dobson, Ann. Mag. N.H. j; 31. Ussuri River, Manchuria. 

Crocidura lasiura lasia Thomas, 1906 

1906. Crocidura leucodon lasia Thomas, Ann. Mag. N.H. /".■ 416. Scalita, near 
Trebizond, 700-1,000 m., Asia Minor. Ranges to Transcaucasia. 

Crocidura lasiura thomasi Sowerby, 191 7 

1917. Crocidura thomasi Sowerby, Ann. Mag. N.H. 20: 318. Mingyong, no miles 
south-east of Seoul, Korea. 

Crocidura lasiura yamashinai Kuroda, 1934 

1934. Crocidura yamashinai Kuroda, J. Mamm. if]: 237. Bampo, Kankyo-hokudo, 

Northern Korea. 
1931. Crocidura lizenkarii Kishida, Zool. Mag. Tokyo, 43: 377, (nom. nud.). 

Crocidura lasiura campuslincolnensis Sowerby, 1945 

1945. Crocidura campus-lincolnensis Sowerby, Musee Heude Notes de Mammalogie, 
No. 3, I. Lincoln Avenue, in the western district of Shanghai, Kiangsu, 
China. (We have not examined this form which from description agrees 
with the larger members of this species.) 


Crocidura olivieri group. (Giant species.) 

Crocidura olivieri Lesson, 1827 Egyptian Giant Shrew 

Approximate distribution of species: Egypt. Perhaps represented in Kenya and 
Abyssinia under the name zcphiri. 

Crocidura olivieri Lesson, 1827 

1827. Sorex olivieri Lesson, Manuel de Mammalogie, 121. Sakkara, Egypt, as 
mummies from catacombs. 

Other Js'amed Forms 

Crocidura fuliginosa Blyth, 1855 

1855. Sorex fuliginosus Blyth, J. Asiat. Sec. Bengal, 24: 362. Schwegyin, near Pegu, 
Burma. The type specimen is in Calcutta, and we have ascertained from the 
curator of the Calcutta Museum that the type specimen has eight upper 
teeth and is therefore a Crocidura. Mrs. Lindsay erroneously transferred 
this form to the genus Suncus. According to Chasen, it occurs in the 
Malay Peninsula. This early name may ultimately have to supersede 
one of the specific names listed above, but we are uncertain of its exact 

Crocidura nicobarica Miller, 1902 

1902. Crocidura nicobarica Miller, Proc. U.S. Nat. Mus. 24: •jyB. Great Nicobar 
Island, Nicobar Islands, Bay of Bengal. Unrepresented in London Based on 
a very large species, head and body 120 mm., tail 90 mm. Basal length of 
skull 26 mm. 

Crocidura andamanensis Miller, 1902 

1902. Crocidura andamanensis Miller, Proc. U.S. Nat. Mus. 24: 777. MacPherson 
Strait, South Andaman Island, Andaman Islands, Bay of Bengal. Un- 
represented in London. Head and body 1 14 mm., tail 86 mm. Basal length 
of skull 24.8 mm. Probably closely allied to nicobarica. 

Crocidura bolfvari Morales Agacino, 1934 

1934. Crocidura bolivari Morales Agacino, Bol. Soc. Esp. H.N. ^4: 93, fig. i. Villa 
Cisneros, Rio de Oro, North-West Africa. We have not examined this form, 
which is likely to be valid unless it represents one of the numerous Ethiopian 
earlier-named species. Condylobasal length 21.6 mm., tail 56 per cent, of 
head and body, from original description. 

Crocidura utsuryoensis Mori, 1937, J. Chosen N.H. Soc. 22: 40, 41. {N.V.) 
Utsuryo Island, off Korea. 

Crocidura nanula Stroganov, 1941, C.R. Acad. Sci. U.R.S.S. 33: 272. Termez, 
Russian Turkestan. According to Vinogradov based on a specimen with the 
dentition of a Crocidura, but may represent an abnormal specimen oi Suncus 


Genus FEROCULUS Kclaart, 1852 

1852. Feroculus Kelaart, Prodr. Faun. Zcylanica, 31. Sorcx macropus BIyth --= Sorex 
feroculus Kelaart. 

I species : Feroculus Jeroculus, page 86 

Feroculus feroculus Kelaart, 1850 Kelaart's Long-clawed Shrew 

Approximate distribution of species: Ceylon. 

P'eroculus feroculus Kelaart, 1850 

1850. Sorex feroculus Kelaart, J. Ceylon Branch Asiat. Soc. :?, 5: 211. Nuwara Eliya, 

6,000 ft., Central Province, Ceylon. 

1851. Sorex macropus Blyth, J. Asiat. Soc. Bengal, 20: 163. Nuwara Eliya, Ceylon. 

1 85 1. Corsira newera-ellia Kelaart, Ann. Mag. N.H. 8: 340. Nuwara Eliya, Cevlon. 
1855. Sorex newera Wagner, in Schreber, Saugeth. Suppl. 5; 564. 

1888. Croeidura macropus Blanford, Fauna Brit. India, Mamm. /; 237. 

Genus SOLISOREX Thomas, 1924 
1924. .S'o&orf.v Thomas, Spolia Zeylan. /j, i: 94. Solisorex pearsoni Th.om3.%. 
I species: Solisorex pearsoni, page 86 

SoUsorex pearsoni Thomas, 1924 Pearson's Long-clawed Shrew 

Approximate distribution of species: Ceylon. 

The presence of two genera of long-clawed shrews in Ceylon and nowhere else is 
disconcerting. But an examination of the characters we have listed above m the key 
to the genera should indicate that Solisorex cannot be referred to Feroculus as a sub- 
genus. Both seem quite distinct from Croeidura or Suncus. 

Solisorex pearsoni Thomas, 1924 

1924. Solisorex pearsoni Thomas, Spolia Zeylan. ij, i: 94, 95. Hakgala, 6,000 ft., 
near Nuwara Eliya, Central Highlands of Ceylon. 

Genus DIPLOMESODON Brandt, 1852 

1852. Diplomesodon Brandt, in Bacr & Helmersen, Beitr. Russ. Reich. ly: 299. 

Sorex pulchellus Li'-htcnstein. 

I species : Diplomesodon pulchellum, page 86 

Diplomesodon pulchellum Lichtenstein, 1823 Piebald Shrew 

Approximate distribution of species: "Sands between the lower Volga and lower 
Emba, the Bolshie Barsuki sands (north of the Sea of Aral), on the north-west coast of 
the Sea of Aral, on Dardsha Peninsula (south-east coast of the Caspian), in Kara- 
kum, Kizil-kum, the sands east of the River Chu and the sandy desert south of Lake 
Balkash between the Rivers Hi and Aksu" (Bobrinskii). 




1823. Sorex pulchellus Lichtenstein, in Eversmann, Reise von Orenburg nach Bok- 
hara, 124. Kirghiz Steppe, Russian Turkestan (collected i May 1821). 


1938. Diplomesodon pukhelhim pallidus Heptner, Bull. Soc. Nat. Moscou, ^j: 165-166. 
Between Merv and Amu Daria, Russian Turkestan. 

Genus ANOUROSOREX Milne-Edwards, 1872 

1870. Anourosorex Milne-Edwards, C.R. Acad. Sci. Paris, yo: 341. (Genus caelebs.) 
1872. Anourosorex Milne-Edwards, Rech. H.N. Mamm. 264. Anourosorex 
squamipes Milne-Edwards. 

1873. Pygmura Anderson, P.Z.S. 229 (footnote). Type not given, but species sub- 
sequently described as Anurosorex assamensis Anderson. 

1875. Anurosorex Anderson, Ann. Mag. N.H. 16: 282. 

I species: Anourosorex squamipes, page 87 

Anourosorex squamipes Milne-Edwards, 1872 Szechuan Burrowing Shrew- 

Approximate distribution of species: China, from Shensi south to Hupeh, Sze- 
chuan, Yunnan; Northern and Western Burma, Assam; Tonkin, in Indo-China. 

Anourosorex squamipes squamipes Milne-Edwards, 1872 

1872. Anourosorex squamipes Milne-Edwards, Rech. H.N. Mamm. 264, pi. 38, fig. i; 

pi. 38A, figs. i-ij. Probably Moupin, Szechuan, China. 
1875. Anourosorex assamensis Anderson, Ann. Mag. N.H. 16: 282. Subsasugu, Assam. 
1923. Anourosorex squamipes capnias G. Allen, Amer. Mus. Nov. No. 100: 10. To-mu- 

lang, Chungtien district, 10,000 ft., Yunnan, China. 
1923. Anourosorex assamensis capito G. Allen, loc. cit. 11. Mucheng, Salween drainage, 

7,000 ft., Yunnan, China. 
Range: Mainland range of the species. 

Anourosorex squamipes yamashinai Kuroda, 1935 

1935. Anourosorex squamipes yamashinai Kuroda, J. Mamm. 16: 288. Taiheizan, 
5,500 ft., Taihokusiu, North Formosa. 

Genus CHIMMAROGALE Anderson, 1877 

1877. Chimmarogale Anderson, J. .\siat. Soc. Bengal, ^6, 2: 262. Crossopus himalavieus 

1 92 1. Crossogale Thomd^s, Ann. Mag. N.H. j: 243. Crossogale sumatrana Thomas, from 

Sumatra (a race of C. phaeura Thomas from Borneo). 

I species in the area covered by this list: 

Chimmarogale platvcephala, page 88 


pal.\earc:tic and Indian mammals i 758-1946 

All named forms are represented in the British Museum. The type oi styani has 
white undcrparts, other specimens are intermediate between this and the normal 
type of dark underparts of the majority of the other specimens, and there is fairly 
clearly only one valid species of this genus in the mainland of Asia, and Japan. The 
na.mei platycephala and himalayica were both published in the year 1842, and it is not 
possible to say exactly which was published first. C. himalayicus was sent to press 
10 October 1842, and according to notes left by J. L. Chaworth-Musters, was pub- 
lished in December 1842 "so probably after plahrephala". We therefore adopt 

Chimmarogale platycephala Temminck, 1842 Himalayan Water-Shrew 

Approximate distribution of species: Japan (Kiushiu), Szechuan, Yunnan, .South- 
Eastern China to Fukien and Chekiang; Laos, Annam, Tonkin (in Indo-China) ; 
Kashmir, Punjab, Sikkim and Northern Burma. 

Chimmarogale platycephala platycephala Temminck, 1842 

1842. Sorex platycephalus Temminck, Fauna Japon. /, Mamm.: 23, plate V, fig. i. 
Near Nagasaki and Bungo, Kiushiu, Japan. Occurs Hondo. 

Chimmarogale platycephal.\ himalayica Gray, 1842 

1842. Crossopiis himalayicus Gray, Ann. Mag. N.H. 10: 261. Chamba, North-Eastern 
Punjab (from notes left by J. L. Chaworth-Musters). Range: recorded from 
Kashmir, Punjab, Sikkim, Darjeeling, Northern Burma, Yunnan (Likiang 
Range), Laos and Tonkin, Indo-China. 

Chimmarogale platycephala styani de \Vinton, 1899 

1899. Chimmarogale styani de \Vinton, P.Z.S. 574. Yangliupa, North-Western 
Szechuan. Has also been recorded from Northern Burma. 

Chimmarogale platycephala leander Thomas, 1902 

1902. Chimmarogale leander Thomas, Ann. Mag. N.H. 10: 165. Kuatun, 1,200 m., 
North-Western Fukien, China. Range includes Chekiang, China. 

Chimmarogale platycephala varennei Thomas, 1927 

1927. Chimmarogale varennei Thomas, P.Z.S. 45. Dakto, Annam, Indo-China. 

Genus NECTOGALE Milne-Edwards, 1870 

1870. .Neclogale Milne-Edwards, C.R. Acad. Sci. Paris, Jo: 341. Nectogale elegans 

I species: Nectogale elegans, page 89 



Nectogale elegans Milne-Edwards, 1870 Szechuan Water-Shrew 

Approximate distribution of species: Sikkim, Bhutan (B.M.), Northern Burma; 
Tibet (B.M.); Szechuan, in China, also recorded from Yunnan and Shensi. 

Nectogale elegans elegans Milne-Edwards, 1870 

1870. Nectogale elegmu Milne-Edwards, C.R. Acad. Sci. Paris, yo: 341. 

Moupin, Szechuan, China. Range: Szechuan, Yunnan, Shensi, Northern 


Nectogale elegans sikhimensis de Winton, 1899 

i8gg. Nectogale sikhimensis de Winton, P.Z.S. 573. Lathong, 10,000 ft., Sikkim. 
Range: Sikkim and Tibet. 


FAMILY: Cynocephalidae, page 89 

Genus: Cynocephalus, page 89 

Thomas (1908) divided the flying lemurs into two genera: Cynocephalus, based on 
Lemur volans Linnaeus, for the Philippine forms which have the first upper incisor 
very reduced and the parietal ridges close together; and Galeopterus, based on 
Galeopithecus temminckii Waterhouse, for the Malayan and East Indies forms which 
have the first upper incisor not so reduced and the parietal ridges widely separated. 
Chasen (1940) follows Thomas, but we prefer to follow Simpson (1945) and include 
all flying lemurs in the genus Cynocephalus. 

Genus CYNOCEPHALUS Boddaert, 1768 

1768. Cynocephalus Boddaert, Dierkundig Mengelwerk 2: 8 (footnote 1). Lemur volans 

1783. Galeopithecus Pallas, Acta Acad. Sci. Petrop. iy8o, i: 208. Lemur volans 

(?) 1840. Galeolemur Lesson, Spec. Mamm. 261. Galeopithecus macrourus Temminck, 

? from Ceylon. (G. macrourus Temminck, 1838, Coup d'GEil Faune lies 

Sonda et Emp. Jap. ix.) This was a flying-squirrel [Petaurista) fide Thomas, 

1908, Ann. Mag. N.H. /.• 252. It is unidentifiable. 
1908. Galeopterus Thomas, Ann. Mag. N.H. /.• 254. Galeopithecus temminckii 

Waterhouse. (G. variegatus temminckii from Sumatra.) Valid as a 


I species in the area covered by this list : 
Cynocephalus variegatus, page 90 



Subgenus GALEOPTERUS Thomas, 1908 

Cynocephalus variegatus Audebert, 1799 Malayan Flyinc; Lemur fCobego or 

Approximate distribution of species: Tenasserim and Southern Indo-China 
(Cochin China), southwards to Malay States, Sumatra, Ja\'a, Borneo and many 
adjacent small islands. 

Cynocephalus variegatus variegatus Audebert, 1799. Extralimital) 
I 799. Galeopithecus variegatus Audebert, H.N. Singes, sig. Rr. Java. 

CvNOCEPH.\Lus varieg.\tus peninsulae Tliomas, 1909 

1909. Galeoplenis peninsulae Thomas, Ann. Mag. N.H. ;?.• 303. Semangko Pass, Malay 

States. Range: Malay Peninsula, Tenasserim. 
Osgood (1932) quoted Galeopterus variegatus subsp. from Cochin-C^hina. 


Special works of reference on this order include: 
Allen, G. M. ic)/^o. Bats. Cambridge, Mass. (Harvard University Press). 
DoBsON, G. E. 1878. Catalogue of Chiroptera in the British Museum. 
Miller, G. S. 1907. The Eamilies and Genera of Bats. Bull. U.S. .hfat. Mus., Xo. 57. 
Andersen, K. 1912. Catalogue of the Chiroptera in the British Museum, i. Megachiroptera. 
(All published.) 

The first is a general work on the order. The second, though old, is still most useful. 
The third is the only comprehensive work on the taxonomy of the order, with keys 
down to genera, and the last is still the only comprehensive work on the Mega- 
chiroptera. Miller seems to recognize too many families, and Simpson (1945, 180) 
takes the view that recent specialists recognize too many genera. Neither of the 
present authors claims any extensive knowledge of this order, which seems very much 
a specialist field. The listing of the species is entirely provisional. Our thanks are due 
to our colleague, Mr. R. W. Hayman, for his help with this order. 

FAMILIES: Emballonuridae, page 103 
Megadermatidae, page 107 
Molossidae, page 132 
Nycteridae, page 106 
Pteropodidae, page 91 
Rhinolophidae, page 109 
Rhinopomatidae, page 10 1 
Vcspcrtilionidae, page 136 

.\nothcr group, the Hipposiderinae, was regarded as a family distinct from the 
Rhinolophidae by Miller, but b>' man\' authors is considered a subfimilv of that 

For ke\s to the various lamilies, sec Miller, 1907, Eamilies and Genera of Rats. 




The classic work on this family is by Knud Andersen (19 12) and it is surprising 
that it is entirely overlooked in the very detailed bibliography given by Simpson, 

1945. P- 273- 

Simpson (p. 54) has attempted some generic reduction in this family, but 
in a rather unfortunate manner; for instance, one genus {"Callinycteris") shown by 
Andersen to be nothing but a synonym oi Eonycteris is listed as valid (p. 55), whereas 
others which are seemingly reasonably distinct are placed as subgenera or in 
the wrong synonymy (for instance, Pterocyon = Eidolon, not Rousettus as listed by 
Simpson) . 

Genera: Cynopterus, page 98 
Eidolon, page 91 
Eonycteris, page 100 
Macroglossus, page 100 
Megaerops, page 99 
Pteropus, page 93 
Rousettus, page 92 
Sphaerias, page 100 

For a key to these genera see Knud Andersen, 1912, Cat. Chiroptera B.M. i 
This author also gives a key to all the species in the present family named before 191 2. 

Subfamily Pteropodinae 

Genus EffiOLON Rafinesque, 1815 

1815. Eidolon Rafinesque, Analyse de la Nature, 54. Vespertilio vampyrus helvus Kerr. 

For note on validity of Eidolon Rafinesque and fixing of type species, see 

K. Andersen, 1908, Ann. Mag. N.H. /.• 432. 
1861. Pterocyon Peters, Mber. Preuss. Akad. Wiss. 423. Pterocyon paleaceus Peters = 

Vespertilio vampyrus helvus Kerr. 

1 88 1. Leiponyx Jentink, Notes Leyden Mus. j.- 60 Leiponyx biittikoferi Jentink = 

Vespertilio vampyrus helvus Kerr. 

1882. Liponyx Forbes, Zool. Record, 18 (for 1881), Mamm. 13. 

I species in the area covered by this list : 
Eidolon sabaeum, page 92 

The first-named species in this genus is Eidolon helvum Kerr, 1792, from Senegal, 
which ranges eastwards to Somaliland, thence southwards as far as the neighbour- 
hood of Cape Town. Andersen separated the Arabian representative as E. sabaeum; 
it is closely allied but is on average a smaller form. 


Eidolon sabaeum K. Andersen, 1907 Arabian Straw-coloured Fruit Bat 

Approximate distribution of species: Southern Arabia. 

Eidolon sabaeum Andersen, 1907 

1907. Pterocvon sabaeus Andersen, Ann. Mag. N.H. ig: 505. Lahej, Aden district, 
Southern Arabia. 

Genus ROUSETTUS Gray, 1821 

1 82 1. Rousettus Gray, London Med. Repository, 15: 299. Plaopus aegyptiacus E. 

1829. Cercopteropus Burnett, Quart. J. Sci. Lit. Art. i: 269. Pteropus aegyptiacus 

1843. Xantharpyia Gray, List Mamm. B.AL xix, 37. Pteropus amplexicaudatus Geoffroy. 

1843. Eleutherura Gray, List Mamm. B.AL xi.x, nom. mid. 

1844. Eleutherura Gray, \'oy. Sulphur, /.• 29. Pteropus leachii Smith, from South 

1852. Cynonveteris Peters, Reise nach Mossambicjue, Saugeth, 25. Pteropus collaris 

Illiger = Pteropus leachii Smith, from South Africa. 
1870. Senonycteris Gray, Cat. Monkeys, Lemurs & Fruiteating Bats, B.\l. 115. 

Pteropus seminudus Kelaart. 

5 species in the area covered by this list: 

Rousettus aegyptiacus, page 92 
Rousettus amplexicaudatus, page 93 
Rousettus arabicus, page 92 
Rousettus leschenaulti, page 93 
Rousettus seminudus, page 93 

A key to these species is given by K. Andersen, 191 2. 

Rousettus aegyptiacus E. Geoffroy, 1810 Egyptian Fruit Bat 

.Approximate distribution of species: Cyprus, Palestine, Syria, Egypt, and 
Ethiopian Africa in part, south to Angola. 

Rousettus aegyptiacus E. Geoflrny, 1810 

1810. Pteropus egyptiacus Geoffroy, Ann. Mus. H.N. Paris, 75.- 96 (misprint), corrected 

to aegyptiacus in 1818, Description de I'Egypte, H.N. :?.• 134, pi. 3, fig. 2. 

Great Pyramid, Giza, Egypt. 
1825. Pteropus geoffroyi Temminck, Mnn. ^L^mm. /.• 197. Senegal, and probably 

north coast of Africa. 

Rousettus arabicus Anderson & dc \Vint(_in, 1902 

.\pproximate distribution of species: Arabia |','\den, Muscat), Kishim Island 
(Persian Gulfj and Karachi. Sind Western India). 



RousETTUS ARABicus Anderson & de \Vinton, 1902 

1902. Rousettus arabicus Anderson & de Winton, Zool. Egypt, Mamm. 86, 88, B9-90. 
Lahej, near Aden, Southern Arabia. 

Rousettus amplexicaudatus E. Geoffrey, 18 10 

Approximate distribution of species: according to Chasen (1940, 29) Northern 
Siam and Tenasserim; Cambodia, Indo-China (K. Andersen); also from Malay 
States, Sumatra, Java, Borneo, Philippine Islands, Timor, Flores, etc. 

Rousettus amplexicaudatus amplexicaudatus Geoffrey, 1810 
1 8 10. Pteropus amplexicaudatus E. Geoffroy, Ann. Mus. H.N. Paris, 75 .• 96, pi. 4. 
Island of Timor. 

Rousettus leschenaulti Desmarest, 1820 

Approximate distribution of species: Kumaon, Nepal, Rajputana, Bhutan Duars, 
Burma, Tenasserim; Peninsula of India (Western Ghats, Bombay, Coorg, etc.); 
North Siam (Chasen, 1940); Tonkin, Indo-China. Has been recorded from Amoy, 
Southern China; Java. 

Rousettus leschenaulti leschenaulti Desmarest, 1820 

1820. Pteropus leschenaulti Desmarest, Encycl. Meth. Mamm. /.• no. Pondicherry, 

1835. Pteropus pyrivorus Hodgson, J. Asiat. Soc. Bengal, ^: 700. Nepal. 
1 84 1. Pteropus pirivarus Hodgson, loc. cit. 10: 908. 

1843. Cynopterus marginalus Gray, List Mamm. B.M. 38. Not of Geoffroy, 1810. 
1843. Cynopterus affinis Gray, loc. cit. 39. Himalayas. 
1870. Eleutherura fuliginosa Gray, Cat. Monkeys, Lemurs 5c Fruiteating Bats, 118. 

Laos Mountains, Siam. 
1870. Eleutherura fusca Gray, loc. cit. 119.? India. 
1873. Cynonycteris infuscata Peters, Mber. Preuss. Akad. \Viss. 487. Calcutta, India. 

Rousettus seminudus Kelaart, 1850 
Distribution: Ceylon. 

Rousettus seminudus Kelaart, 1850 

1850. Pteropus seminudus Kelaart, J. Ceylon Br. Asiatic Soc. 2: 216. Mount Lavinia, 

Genus PTEROPUS Brisson, 1762 

1762. Pteropus Brisson, Regn. Anim. 13, 153-155. Pteropus niger (Kerr). Hopwood 
(1947) would ignore Brisson and date Pteropus from Erxleben, 1777, Syst. 
Anim. 130, with the same type species. 

1799. Spectrum Lacepede, Tabl. Mamm. 15. Pteropus niger (Kerr). Not of Scopoli, 

1866. Eunycteris Gray, P.Z.S. 64. Pteropus phaiops Temminck = Pteropus melanopogon 
Peters, from Amboina. 



Pteropus [contd.] 

1870. Pselaphon Gray, Cat. Monkeys, Lemurs & Fruiteating Bats, B.M. 1 10. Pteropus 

pselaphon Layard, from Bonin Islands. 
1899. Sericonycteris Matschie, Megachiroptera Berlin Mus. 6, 30. Pteropus subniger 

(Kerr) from Reunion and Mauritius. 
1907. Desmalopex Miller, Fam. & Gen. Bats, 60. Pteropus leucopterus Temminck, from 

Luzon, Philippine Islands. 

The genus appears to need revision ; in the present region^ the following si.x species 
seem most likely to prove valid : 

Pteropus dasymallus, page 94 
Pteropus hypomelanus, page 95 
Pteropus Ivlei, page 96 
Pteropus mariannus, page 95 
Pteropus melanotus, page 96 
Pteropus vampyrus, page 96 

Andersen di\ided this large genus into 17 species groups and 82 species. Four of 
his groups occur in the present region. 

Pteropus subniger group 

(This is the "hypomelanus group" of Andersen, but P. subniger (Kerr, 1792), from 
Reunion Island, east of Madagascar, is the prior name, and we feel that species 
groups should be named after the earliest-named species which they contain.) 

Pteropus dasymallus Temminck, 1825 Liukiu Islands Flying Fox 

Approximate distribution of species: Liukiu Islands and Formosa. 

Pteropus dasymallus dasymallus Temminck, 1825 

1824. Pteropus rubrkollis Siebold, de Hist. Nat. Jap. 13. Liukiu Islands (Andersen, 

1912). Not of Geoffrey, 1810. 

1825. Pteropus dasymallus Temminck, Mon. Mamm. /.• 180, pi. 10. Type locality 

restricted to Kuchino-Erabu, North Liukm Islands (Kuroda, 1933). 
1929. Pteropus vamagatai Kishida, Lansania, Tokyo, /, 8: 125. Kuchino-Erabu, 
North Liukiu Islands. 

Pteropus dasymallus formosus Sclater, 1873 

1873. Pteropus formosus Sclater, P.Z..S. 193, pi. 22. Taku, Formosa. 

Pteropus dasymallus inopinatus Kuroda, 1933 

1933. Pteropus dasymallus inopinatus Kuroda, J. Mamm. 14: 314. Nago-Mura, Kunjan, 
Okinawa Island, Liukiu Islands. 

Pteropus dasymallus yayeya.mae Kuroda, 1933 

1933. Pteropus dasymallus yayeyamae Kuroda, J. Mamm. 14: 315. Ishigaki, Yayeyama 
group, South Liukiu Islands. 



Pteropus hypomelanus Temminck, 1853 Small Flying Fox 

Approximate distribution of species : Cochin-China, Siam, Mergui Archipelago, 
islands off Lower Siam, Straits of Malacca, islands west of Sumatra, Natuna and 
Anamba Islands, islands off Borneo; Celebes, Philippine Islands, New Guinea. 

(Pteropus hypomelanus hypomelanus Temminck, 1853. Extralimital) 
1853. Pteropus hypomelanus Temminck, Esq. Zool. Cote Guine, 61. Ternate Island 
(Gilolo group, between Celebes and New Guinea). 

Pteropus hypomelanus condorensis Peters, 1869 

1869. Pteropus condorensis Peters, Mber. Preuss. Akad. \Viss. 393. Pulau Condor 

(Condor Island), off Cambodia, Indo-China. Range: said to occur Cam- 
bodia and Siam (Andersen). 

Pteropus hypomelanus geminorum Miller, 1903 

1903. Pteropus geminorum Miller, Smith's Misc. Coll. 4^: 60. South Twin Island, 

Mergui Archipelago. Range includes certain West Siamese Islands (see 

Chasen, 1940, Bull. Raffles Mus. /j.- 22). 

Pteropus (?) hypomelanus satyrus Andersen, 1908 

1908. Pteropus satyrus Andersen, Ann. Mag. N.H. 2: 362. Narcondam, North Anda- 
man Islands, Bay of Bengal. (Andersen retained this form as a species, but 
from descriptions it seems very close to hypomelanus.) 

Andersen also referred the following to the present group : 

Pteropus faunulus Miller, 1902 

1902. Pteropus faunulus Miller, Proc. U.S. Nat. Mus. 24: 785. Car Nicobar, Nicobar 
Islands, Bay of Bengal. 

Pteropus mariannus group 

Pteropus mariannus Desmarest, 1822 

Approximate distribution of species: described from Mariana Islands, Western 
Pacific; represented in the Liukiu Islands. 

(Pteropus mariannus mariannus Desmarest, 1822. Extralimital) 
1822. Pteropus mariannus Desmarest, Encycl. Meth. (Mamm.) 2: 547. Mariana 
Islands, Western Pacific. 

Pteropus mariannus loochoensis Gray, 1870 

1870. Pteropus loochoensis Gray, Cat. Monkeys, Lemurs & Fruiteating Bats, B.M. 106. 

Liukiu Islands. 
1892. Pselaphon luchuensis Seitz, Mitt. Dtsch. Ges. Naturk. Ostasiens, 5; 364. {N.V.) 
1894. Pteropus keraudreni var. loochooensis Fritze, Zool. Jb. Syst. y: 854. Okinawa, 

Liukiu Islands. 



Pteropus mclanotus group 

Pteropus melanotus Blyth, 1863 Nicobar Flying Fox 

Approximate distribution of species: Nicobar Islands, and as here understood, 
Andaman Islands; Xias and Engano Islands; \Vestern Sumatra; Christmas Island 
(south of Java). 

Pteropus melanotus melanotus Blyth, 1863 

1846. Pteropus cdulis Blyth, J. Asiat. Soc. Bengal, /j; 367. Not of E. Geoflroy, 1810. 
1861. Pteropus mcohancus Fitzinger, S.B. Akad. Wiss. Wien. 42: 389, nom. nud. 
1863. Pteropus melanotus Blyth, Cat. Mamm. Mus. Asiat. Soc. 20. Nicobar Islands, 
Bay of Bengal. 

Pteropus (?) melanotus tytleri Mason, 1908 

1908. Pteropus trtlerl Mason, Rec. Ind. Mus. 2: 162. Rutland Island, South Andaman 
Islands, Bay of Bengal. 
On the status of this form, see K. Andersen, 1912, Cat. Chiropt. i: 821. It does 
not seem, from present knowledge, that this form should be granted the rank of a 
valid species. According to Chasen, the other members of Andersen's melanotus 
group, respectively from Nias Island and Engano Islands, west of Sumatra, and 
Christmas Island, south of Java, are all races of P. melanotus, although .Andersen 
listed them all binominally. 

Pteropus vampyrus group 

\Ve provisionally follow Andersen in listing members of the vampyrus group as 
species, though with the exception of P. lylei it seems more likely, as Andersen himself 
suggests on p. 32'-,, that they are in reality all members of one species for which the 
first name is P. vampyrus. 

Pteropus lylei K. Andersen, 1908 

Approximate distribution of species: Siam (Bangkok region) and Saigon, Cochin 

Pteropus lylei Andersen, 1908 

1908. Pteropus lylei Andersen, Ann. Mag. N.H. 2: 367. Bangkok, Siam. 

Pteropus vampyrus Linnaeus, 1758 Malayan Large Flying Fox 

Approximate distribution of species: has been recorded from Tenasserim (^ool. 
Record, 1926, Mamm. 47); Annam, and Phu Q_uoc Island, Indo-China (Osgood, 
1932) Also from Malay States, Sumatra, Java, Borneo, Philippine Islands, Bali, 
Timor, and numerous adjacent small Malaysian islands. 

(Pteropus vampvrus vampyrus Linnaeus, 1758. Extralimital) 
1758. Vespertilio vampyrus Linnaeus, Syst. Nat. loth ed. /.• 31. Java. 



Pteropus vampyrus malaccensis Andersen, 1908 

1908. Pteropus vampyrus malaccensis Andersen, Ann. Mag. N.H. 2: 368. Kuala Tem- 
beling, Pahang, 200 ft., Malay Peninsula. Range: Malay States, Sumatra, 
some adjacent islands; northwards to Indo-China, as noted above, and 

Pteropus giganteus Briinnich, 1782 Indian Flying Fox 

Approximate distribution of species: Ceylon, Peninsula of India (widely distri- 
buted), northwards to Rajputana, Cutch, Kathiawar and district, Kumaon, Punjab; 
Nepal, Sikkim, Bhutan Duars, Assam, Manipur, Pegu in Burma. 

Pteropus giganteus giganteus Brunnich, 1782 

1782. Vespertilio gigantea Brunnich, Dyrenes Historic, /.• 45. Bengal, India. 

1825. Pteropus medius Temminck, Mon. Mamm. /.• 176. Calcutta; Pondicherry, India. 

1828. Pteropus edwardsi I. Geoffroy, Diet. Class. H.N. i^: 699. Bengal. Not of E. 

Geoffroy, 18 10. 
1870. Pteropus kelaarti Gray, Cat. Monkeys, Lemurs & Fruiteating Bats, B.M. 104. 


Range: Ceylon, Peninsular India north to Punjab, and apparently eastwards to 
Sikkim, Bhutan Duars, Pegu (Wroughton, 1918). 

Pteropus giganteus leucocephalus Hodgson, 1835 

1835. Pteropus leucocephalus Hodgson, J. Asiat. Soc. Bengal, 4: 700. Central region of 

1839. Pteropus assamensis M'Clelland, P.Z.S. 148. Assam. 

1840. Pteropus rubricollis (misspelt rubicollis) Ogilby, Madras J. Lit. 12: 146. Assam. 

Nom. nud. Not of E. Geoffroy, 1810. 

Range: Nepal, Assam, Manipur. 

Other forms listed as species by Andersen in the present group : 

Pteropus ariel G. Allen, 1908 

1908. Pteropus ariel G. Allen, Bull. Mus. Comp. Zool. Harv. 52, 3: 28, 

fig. I. Male Atoll, Maldive Islands (south-west of Southern Peninsular 


Pteropus intermedius Andersen, 1908 

1908. Pteropus intermedius Andersen, Ann. Mag. N.H. 2: 368. Amherst, Tenasserim. 
(This seems intermediate between P. vampyrus and P. giganteus, 
suggesting that all these forms are really probably only subspecies of 
P. vampyrus.) 

Incertae sedis 

Pteropus daitoensis Kuroda, 1921, J. Mamm. 2: 210. Kita-Daitojima, Daito Islands, 
South-Eastern Liukiu Islands. 



Genus CYNOPTERUS F. Cuvier, 1824 

1824. Cynopterus F. Cuvier, Dents Mamm. 248. Pteropus marginatus Geoffroy = Vesper- 

tilio sphinx Vahl. 
1828. Pachvsoma E. Geoffroy, Cours. H.N. Mamm. 13, le^on 26. Not of Macleay, 

1906. Niadius Miller, Proc. Biol. Soc. Washington, ig: 83. Cynopterus pnnceps Miller, 

from Nias Island, Western Sumatra. 

2 species in the area covered by this list : 
Cynopterus brachyotis, page 98 
Cynopterus sphinx, page g8 

These two species are closely allied, but occur together. Other species occur in the 
Malay region. 

Cynopterus sphinx Vahl, 1797 Short-nosed Fruit Bat 

Approximate distribution of species: Hainan; Peninsula of India, where it is 
widely distributed, Ceylon, Bengal, Kumaon, Sikkim, Bhutan Duars, Burma, Indo- 
China, Siam, also Sumatra, Java, Bali, Lombok, Timor. 

Cynopterus sphinx sphinx Vahl, 1797 

1797. Vespertilio sphinx Vahl, Skr. Nat. Selsk Copenhagen, 4, i: 123. Tranquebar, 

Madras, India. 
1797. Vespertilio fibulatus Vahl, loc. cit. 124. Tranquebar, Madras, India. 
1803. Pteropus pusillus E. Geoffroy, Cat. Mamm. Mus. H.N. 49. India. Not valid, as 

according to Sherborn this was never published. 
1810. Pteropus marginatus E. Geoffroy, Ann. Mus. H.N. Paris, 75.- 97, pi. v. Bengal. 

1837. Pachvsoma brevicaudatum Temminck, Mon. Mamm. 2: 92. Calcutta, India. 
1870. Cynopterus marginatus var. ellioti Gray, Cat. Monkeys, Lemurs & Fruiteating 

Bats, B.M. 122. 
Range: Ceylon, Peninsula of India, Kumaon, Sikkim, Bhutan Duars, Sylhct 
(Assam), Chin Hills and Shan States, Burma, Northern Siam. 

Cynopterus sphinx gangeticus Andersen, 1910 

1 910. Cynopterus sphinx gangeticus Andersen, Ann. Mag. N.H. 6: 623. Lucknow, 

United Provinces, India. Range includes Central Provinces and Palanpur, 


Cynopterus brachyotis Miiller, 1838 

Approximate distribution of species: has been recorded from near Canton, 
Southern China; Ceylon; Andaman and Nicobar Islands; Tenasserim, Burma, 
Assam; Siam; Malay States, Sumatra, Java, Borneo, and adjacent small islands, 
Celebes, Philippine Islands. 

(Cynopterus brachyotis brachyotis Miiller, 1838. Extralimital) 

1838. Pachvsoma brachyotis Miiller, Tijdschr. Natuur. Gesch. 5, i: 146. Borneo. 

Range: Lower Siam, east to Celebes, Philippines. 



Cynopterus brachyotis scherzeri Zelebor, 1869 

1869. Cynopterus marginatus var. {Pachysoma scherzeri) Zelebor, Reise Novara, 


13. Car Nicobar, Nicobar Islands. Range includes Great Nicobar Island. 

Cynopterus brachyotis ceylonensis Gray, 1870 

1870. Cynopterus marginatus var. ceylonensis Gray, Cat. Monkeys, Lemurs & Fruit- 

eating Bats B.M. 122. Ceylon. 

Cynopterus brachyotis brachysoma Dobson, 1871 

187 1. Cynopterus brachysoma Dobson, Proc. Asiat. Soc. Bengal, 105. Andaman Islands, 

Bay of Bengal. 
1873. Cynopterus marginatus var. andamanensis Dobson, loc. cit. 148, nom. nud. J. Asiat. 
Soc. Bengal, 42: 201, pi. xiv, fig. 5. 

Cynopterus brachyotis angulatus Miller, 1898 

1898. Cynopterus angulatus Miller, Proc. Acad. Nat. Sci. Philadelphia, 316. 
Trang, Lower Siam. Range: Kindat (Chindwin), Western Burma, 
Tenasserim, Siam (Nan, Bangkok, Chiengmai, etc.), Cambodia and 
Annam, Natuna Islands and Anamba Islands, various small islands off 

Cynopterus brachyotis hoffeti Bourret, 1944 

1944. Cynopterus brachyotis hoffeti Bourret, Notes Trav. Ecole Sup. Sci: Hanoi, j: 4. 
Cho-Bo, near Hanoi, Tonkin, Indo-China. 

Genus MEGAEROPS Peters, 1865 

1841. Megera Temminck, Mon. Mamm. 2: 274. Pachysoma ecaudatum Temminck. 
1 84 1. Megaera Temminck, loc. cit. 359. Not of Wagler, 1830, or Robineau-Devoidy, 

1865. Megaerops Peters, Mber. Preuss. Akad. Wiss. 256. Megaera ecaudata Temminck. 

Osgood has recorded this principally Malaysian genus from Indo-China. Simpson 
(1945) would refer it to Ptenochirus, Peters, 1861, from which it seems reasonably 

I species : Megaerops ecaudatus, page 99 

Megaerops ecaudatus Temminck, 1837 Temminck's Fruit Bat 

Approximate distribution of species: Malay States, Sumatra, Borneo; recorded 
from Annam, in Indo-China, by Osgood (1932). 

Megaerops ecaudatus Temminck, 1837 

1837. Pachysoma ecaudatum Temminck, Mon. Mamm. 2: 94. Padang, Western 



Genus SPHAERIAS Miller, 1906 

1906. Sphaerias Miller, Proc. Biol. Soc. Washington, ig: 83. Cynoplerus hlanfordi 

I species: Sphaerias hlanfordi, page 100 

Sphaerias blanfordl Thomas, iSgr Blanford's Fruit Bat 

.\ppniximate distribution of species: Karin Hills, Burma, and Siam according to 

Sphaerias blanfordi Thomas, 1891 

1891. CynopUrus hlanfordi Thomas, Ann. Mus. Stor. Nat. Geneva, i>, 10: 884, 921- 
922, pi. XI, figs. 1-2. Lcito, Cheba, Karin Hills, 1,000 m., Burma. 

Subfamily Macroglossinae 

Genus EONYCTERIS Dobson, 1873 

1873. Eonycteris Dobson, Proc. Asiat. Soc. Bengal, 148. Alacroglossus spelaeus Dobson. 
1889. Callinvcteris ^entmk, Notes Lcyden Mus. //.• 209. Callinycteris rosenbergii ]cn- 
tink, from Clclebes. 

I species in the area covered by this list: 
Eonycteris spelaea, page 1 00 

Eonycteris spelaea Dobson, 1871 Dobson's Long-tongued Fruit Bat 

Approximate distribution of species: Burma, Indo-China, Siam, Malay States, 
Sumatra, Java, Borneo, Luzon (Philippine Islands). 

Eonycteris spelaea Dobson, 1871 

1871. Macroglossiis spelaeus Dobson, Proc. Asiat. Soc. Bengal, 105, 106. Farm Caves, 
Moulmein, Tenasserim. Range includes Nan in Siam, Tonkin, Laos, 
Cochin-China in Indo-China, Malay States, Sumatra, Java, Borneo. 

Genus MACROGLOSSUS F. Cuvier, 1824 

1824. Macrnglossiis C'uvier, Dents Mamm. 248. Pternpus minimus Geoffroy. 

1840. Kiodolus Blyth, in Cuvier, Anim. Kingd. 69. New name for Maeroalossus, under 

the impression that it was preoccupied by Maeroglossum Scopuli, 1777. 
1848. Rhynchocyon Gistel, Naturg. Thierr. ix. Not of Peters, 1847. 
1 89 1. Carporvtctcris Lydekker, in Flower & Lydekkcr, Mamm. Living & Extinct, 654. 

New name for Macroglossiis Cuvier. 
1902. Odontonycteris ]cn\.mk. Notes Leyden Mus. 2^: 140. Odonto/mleris wnr// Jentink 

= Macroglossiis lagochiliis Matschie, from Burn, Moluccas. 

I species in the area cosercd by this list : 
Macroglossiis minimus, page loi 



Macroglossus minimus E. Geoffroy, 1810 Small Long-tongued Fruit Bat 

Approximate distribution of species: Tenasserim; Malay States, Sumatra, Java, 
Bali, and a few adjacent islands. (Tate also quotes it from Darjeeling.) 

(Macroglossus minimus minimus Geoffroy, 1810. Extralimital) 

1810. Pleropus minimus E. Geoffroy, Ann. Mus. N.H. Paris, i^: 97. Java. 

Macroglossus minimus sobrinus Andersen, igii 

191 1. Macroglossus minimus sobrinus Andersen, Ann. Mag. N.H. y: 642. Gunong Igari, 
Perak, 2,000 ft., Malay States. Range: northwards to Tenasserim. 

sub-order Microchiroptera 

Genus: Rhinopoma, page loi 

Genus RHINOPOMA Geoffroy, 1818 

1818. Rhinopoma Geoffroy, Description de I'Egypte, 2: 113. Vesperlilio microphyllus 

1 82 1. Rhynopoma Bowdich, Anal. Nat. Class. Mamm. 30. Vesperlilio microphyllus 


3 species in the area covered by this list: 
Rhinopoma hardwickei, page 102 
Rhinopoma kinneari, page 102 
Rhinopoma microphjllum, page 102 

Formerly, as by Dobson and Blanford, all known forms were referred to a single 
species, R. microphyllum, although Dobson stated that the Asiatic representatives 
differed in certain respects from the African ones. See particularly Thomas, 1903, 
Ann. Mag. N.H. 11: 496, and WYoughton, 1912, J. Bombay N.H. Soc. 21: 767. 
Wroughton gave a key to the known forms. Thomas, in erecting cystops, apparently 
failed to compare it with hardwickei. It seems clear that in Egypt there are two species 
(a larger and a smaller) occurring together. It also seems clear from W'roughton's key 
that there are two groups of species, a larger (rare) group and a smaller group (or 
species) which occurs throughout much of the range of the genus, at least as far as 
this list is concerned. The prior name for the smaller species is R. hardwickei. Accord- 
ing to Wroughton, this and its allies differ from the large microphyllum group both in 
an external and in a cranial character, but it is very difficult to believe that there are 
in reality four distinct species of smaller Rhinopoma, and the smaller named species 
are here provisionally made representative races of the first-named hardwickei. The 
large Indian R. kinneari is, from descriptions, larger than the Egyptian R. micro- 
phyllum, and widely separated from it geographically. Another equally large species 
has been described from Sumatra. 



Rhinopoma microphyllum Brunni-ch, 1782 Larger Rat-tailed Bat 

Approximate distribution of species: Arabia, Egypt, Palestine, perhaps Persia. 

Rhinopoma microphyllum Brunnich, 1782 

1782. Vesperlilio microphyllus Brunnich, Dyrenes Hist. /.• 50, pi. 6, figs. 1-4. Arabia 
and Egypt. 

Rhinopoma kinneari \Vroughton, 1912 

Approximate distribution of species: Cutch, Kathiawar, Nimar and Bengal, India. 

Rhinopoma kinneari VVroughton, 19 12 

1 91 2. Rhinopoma kinneari VVroughton, J. Bombay N.H. Soc. 21, 3: 767. Bhuj, Cutch, 


Rhinopoma hardwickei Gray, 1831 Lesser Rat-tailed Bat 

.Approximate distribution of species, as here understood: Peninsular India, known 
from Rajputana, Allahabad, Khandesh, Dharwar, Sind, Cutch, Palanpur, Kathia- 
war, Gwalior, Central Provinces, Bellary, Bengal; (Kashmir (Dobson) and Burma 
(Blanford) ) ; Lower Siam; Arabia, Palestine and Persia; Egypt and the Sudan, west 
to Asben region, south to Lake Rudolf. 

Rhinopoma hardwickei hardwickei Gray, 1831 

1 83 1. Rhinopoma hardwickii Gray, Zool. Misc. 37. India. Range: Indian range of 
species above, and Lower Siani. 

Rhinopom.\ hardwickei cystops Thomas, 1903 

1903. Rhinopoma cystops Thomas, Ann. Mag. N.H. //.■ 496. Luxor, Egypt. Range: 
EgN'pt and Sudan, westwards to Asben. 

Rhinopoma hardwickei muscatellum Thomas, 1903 

1903. Rhinopoma muscatellum Thomas, Ann. Mag. N.H. //.• 498. Wadi Bani Ruha, 
Muscat, Arabia. 

Rhinopoma hardwickei arabium Thomas, 19 13 

1913. Rhinopoma cystops arabium Thomas, Ann. Mag. N.H. 12: 89. W'asil, Yemen, 

4,000 ft., Arabia. R^nge: to Midian (North-Western Arabia) and Palestine. 

Rhinopoma hardwickei seianum Thomas, 191 3 

191 3. Rhinopoma muscatellum seianum Thomas, Ann. Mag. N.H. 12: 90. Seistan, Persia. 

Rhinopoma hardwickei pusillum Thomas, 1920 

1920. Rhinopoma pusillum Thomas, J. Bombay N.H. Soc. 2y: 25. Sib, South-Eastern 




Genera: Coleura, page 103 

Emballonura, page 103 
Taphozous, page 104 

A key to these, and all genera of Microchiroptera, will be found in Miller, 1907, 
Families & Genera of Bats (Emballonuridae key, p. 85). 

Genus EMBALLONURA Temminck, 1838 

1838. Emballonura Temminck, Tijdschr. Natuur. Gesch. 5; 22. Emballonura monticola 

I species in the area covered by this list: 
Emballonura monticola, page 103 

Emballonura monticola Temminck, 1838 Sheath-tailed Bat 

Approximate distribution of species: Tenasserim, Northern Siam (Bangkok, 
quoted by Chasen, 1940). Malay States, Java, Sumatra, Borneo, and certain ad- 
jacent small islands. Philippine Islands, according to Dobson. 

Emballonura monticola Temminck, 1838 

1838. Emballonura monticola Temminck, Tijdschr. Natuur. Gesch. ^: 25, pi. ii, figs. 

1-2. Java. 
(?) 1891. Emballonura semicaudata Blanford, Fauna Brit. India, Mamm. 2: 345. ? Not 

ofPeale, 1848. 
1898. Emballonura peninsularis Miller, Free. Acad. Nat. Sci. Philadelphia, 323. Trang, 

Lower Siam. 
Range: as above. 

Genus COLEURA Peters, 1867 
1867. Coleura Peters, Mber. Preuss. Akad. Wiss. 479. Emballonura afra Peters. 
I species in the area covered by this list : 
Coleura gallarum, page 103 

The first-named species in this genus is C. afra Peters, 1852, from Portuguese East 
Africa. The South Arabian form is very like it apparently, but from descriptions is 
a little smaller in forearm and upper toothrow measurements. 

Coleura gallarum Thomas, 1915 Aden Sheath-tailed Bat 

Approximate distribution of species: Somaliland, Sudan and Congo, to Aden 
district. Southern Arabia. 

Coleura gallarum gallarum Thomas, 1915 

1915. Coleura gallarum Thomas, Ann. Mag. N.H. i^: 576. Zeyla, British Somaliland. 
Ranges to Aden district. South- Western Arabia. 



Genus TAPHOZOUS GcofTroy, 1818 

t8i8. Taphoznus Geoffroy, Description de I'Egypte, 2: 113. Taphozous perfnratus 

1838. Saccnlaimus Tcniminrk, Tijclscln'. Natuur. Gesch. 5: 6. Ttiphnzoi/s snccninimus 

1866. Saccolaimus Gray, Ann. Mag. N.H. ly: 92. Taphozous saccolaimus Teniminck. 

Valid as a subgenus. 
1876. Taphonvcleris V)oh%on, P.Z.S. i8j'^: 548. Taphozous saccolaimus Tcrammck. 
1922. Liponrcferis Thomas, Ann. Mag. N.H. g: 267. Taphozous nudiventris Cretzschmar. 

Valid as a subgenus. 

7 species in the area covered by this list : 
Taphozous kach'hensis, page 106 
Taphozous longimamis, page 104 
Taphozous melanopogon, page 105 
Taphozous nudiventris, page 105 
Taphozous perforatus, page 104 
Taphozous saccolaimus, page 106 
Taphozous theobaldi, page 105 

\Ve agree with Simpson that Saccolaimus and Liponycteris, often given generic rank, 
mav well be regarded as subgenera. Miller, in his Families & Genera of Bats, referred 
all these groups to a single genus, and Tate, 1941, Amer. Mus. Nov. No. 1141 : i, in a 
review of the Eastern members of the genus, seems to come to the same conclusion. 
Dobson ''1878, 379) gives a key to the species. 

Subgenus TAPHOZOUS Geoffroy, 1818 

Taphozous perforatus E. Geoffroy, 1818 Tomb Bat 

Approximate distribution of species: Egypt, southwards to Sudan and Kenya; 
Arabia; Cutch and Kathiawar, in India. 

Taphozous perforatus perforatus E. Geoffroy, 1818 

1818. Taphozous perforatus Geoffroy, Description de I'Egypte, 2: 126. Egypt. Range: 
also listed from Cutch and Kathiawar, India, by \Vroughton (1918). 

T.\PHOzous PERFORATUS Hi^EDiNus Thomas, 1915 

1915. Taphozous perforatus haedinus Thomas, J. Bombay N.H. Soc. 24: 62. Chanler 

Falls, Northern Guaso Nyiro, Kenya, East Africa. Range: to Aden, 

Southern Arabia, and district. 

Taphozous longimanus Hardwicke, 1825 

Approximate distribution of species: Ceylon, Peninsula of India, where it appears 
to be quite widely distributed, northwards to Palanpur, Bengal, thence to Burma, 
Tenasserim, Malay States, Sumatra, Java, Borneo, probably Flores, whence Dobson 
described a variety. 



Taphozous longimanus longimanus Hardwicke, 1825 

1825. Taphozous longimanus Hardwicke, Trans. Linn. Soc. London, i^: 525. Calcutta, 

Bengal, India. 
1841. Taphozous fulvidus Blyth, J. Asiat. Soc. Bengal, 10: 975. Darjeeling, North- 

Eastern India. 

1 84 1. Taphozous brevicaudus Blyth, loc. cit. 976. Travancore, India. 

1842. Taphozous canton Blyth, loc cit. 11: 784. Calcutta, India. 
Range: Indian range, as listed above. 

Taphozous tnelanopogon Temminck, 1841 Black-bearded Tomb Bat 

Approximate distribution of species: Java, Malay States, Sumatra, Borneo (prob- 
ably represented in Philippine Islands), Laos, in Indo-China, Tenasserim, Burma, 
also widely distributed in Peninsula of India, south at least to Western Ghats; 
Yunnan, China. 

Taphozous melanopogon melanopogon Temminck, 1841 

1 84 1. Taphozous melanopogon Temminck, Mon. Mamm. 2: 287. Bantam, Western 
Java. Range: Java, also Indian localities as above, Yunnan and Laos. 

1841. Taphozous bicolor Temminck, loc. cit. 290. India. 

(?) 1913. Taphozous solifer Hollister, Proc. Biol. Soc. Washington, 26: 157. Thought 
to be from Pekin, Chihli, China. See G. Allen, 1938, Mamm. China & 
Mongolia, /.■ 160, for a note on this form. Allen thought there was a mistake 
in the locality and that it probably came from some more tropical locality, 
perhaps the Philippines. It was said to be very close to T. philippinensis, 
Waterhouse, 1845, which probably represents melanopogon. 

Taphozous theobaldi Dobson, 1872 

Approximate distribution of species: Tenasserim; Nimar (Central Provinces 
district, India); Indo-China (Bourret, 1944); Malay States; Java. 

Taphozous theobaldi theobaldi Dobson, 1872 

1872. Taphozous theobaldi Dobson, Proc. Asiat. Soc. Bengal, 152. Tenasserim. 

Taphozous theobaldi secatus Thomas, 1915 

1915. Taphozous theobaldi secatus Thomas, J. Bombay N.H. Soc. 24: 60. Asirgarh, 
Nimar, Central Provinces, India. 

Subgenus LIPONl'CTERIS Thomas, 1922 

Taphozous nudiventris Cretzschmar, 1830 vel 1831 Naked-bellied Tomb Bat 
Approximate chstribution of species: Palestine., Arabia; Egypt; Sudan. 

Taphozous nudfventris Cretzschmar, 1830 vel 1831 

1830 vel 1 83 1. Taphozous nudiventris Cretzschmar in Riippell, Atlas Reise Nordl. 

Afrika, Saugeth. 70, fig. 27b. Giza, Egypt. 
1841. Taphozous nudiventer Temminck, Mon. Mamm. 2: 280. 



Taphozous kachhensis Dobson, 1872 

Approximate distribution of species: India, from Sind, Cutch, Palanpur, Kathia- 
war, also parts of the Peninsula (Bellary, Mysore, Khandesh) ; Bengal and Sikkim; 
Burma; Malay States; Iraq. 

Taphozous kachhensis kachhensis Dobson, 1872 

1872. Taphozous kachhensis Dobson, J. Asiat. Soc. Bengal, 41, 2: 221. Cutch, India. 
Range: Indian range, as above, excluding Burma. 

Taphozous kachhensis magnus Wettstein, 191 3 

1913. Taphozous magnus Wettstein, Ann. Naturh. (Mus.) Hofmus. Wien, 2y: 466, 

pi. XX, figs. 1-6. Basra, Euphrates, Iraq. 
191 5. Taphozous kachhensis babylonicus Thomas, J. Bombay N.H. Soc. 24: 58. 

Euphrates River, Iraq. 

Taphozous kachhensis nudaster Thomas, 19 15 

191 5. Taphozous kachhensis nudaster Thomas, J. Bombay N.H. Soc. 24: 59. Pagan, near 
Mt. Popa, Burma. 

Subgenus SACCOLAIMUS Lesson, 1842 

Taphozous saccolaimus Temminck,- 1838 Pouch-bearing Bat 

Approximate distribution of species: Ceylon, Peninsula of India, to Bengal, 
perhaps Burma; Malay States, Sumatra, Java. 

(T.^PHOzous SACCOLAIMUS s.^ccoLAi.MUs Tcmmiuck, 1838. Extralimital) 
1838. Taphozous saccolaimus Temminck, Tijdschr. Natuur. Gesch. j.' 14. Java. 

Taphozous saccolaimus crassus Blyth, 1844 

1844. Taphozous crassus Blyth, J. Asiat. Soc. Bengal, ij: 491. Mirzapore, Allahabad, 

United Provinces, India. 
(?) 1844 Taphozous pulcher Blyth, J. Asiat. Soc. Bengal, /jj; 492. Madras, India. 
Range: Mainland range as above, and Sumatra. 


Genus: Nycteris, page 106 

Genus NYCTERIS Cuvier & Geoffroy, 1 795 

1795. Nycteris Cuvier c& Geoffroy, Mag. Encyclop. 2: 186, nom. nud. Vespertilio 
hispidus Schreber. Name validated by Opinion iii of International Com- 
mission on Zoological Nomenclature. 

1803. Nicteris Desmarest, Nouv. Diet. H.N. i§: 501. 

1838. Petalia Gray, Mag. Zool. Bot. 2: 494. Nycteris javanicus Geoffroy. 

1866. Nyclerops Gray, P.Z.S. 83. Nycterops pilosa Gray = Vespertilio hispidus Schreber. 



2 species in the area covered by this Hst: 
Nycteris javanica, page 107 
Nycteris thebaica, page 107 

On this genus see Andersen, 1912, Ann. Mag. N.H. 10: 546; Dobson, 1878, Cat. 
Chiroptera B.M. 162 (key to species); Tate, 1941, Amer. Mus. Nov. No. 1140, 7. 

The first named species in this genus is apparently M. hispida Schreber, 1774, from 
Senegal, which is described as having relatively shorter ears than the two species 
which come into the region now under discussion. Dobson distinguishes these two 
principally by the fact thai in N.javanica the second lower premolar is two-thirds the 
size of the first and lies in the toothrow, whereas in N. thebaica the tooth is minute, 
and is internal to the toothrow; and by the shape of the tragus. 

Nycteris javanica Geoffroy, 1813 Javan Slit-faced Bat 

Approximate distribution of species: Tenasserim, Malay States, Java, Borneo, 

(Nycteris javanica jav.anica E. Geoffroy, 181 3. Extralimital) 
1813. Nycteris javanicus Geoffroy, Ann. Mus. N.H. Paris, 20: 20. Java. 

Nycteris javanica tragata Andersen, 191 2 

igi2. Petalia tragata Andersen, Ann. Mag. N.H. 10: 546. Bidi Caves, Sarawak, 
Borneo. Range includes Malay States and Tenasserim. 

Nycteris thebaica Geoffroy, 18 18 Egyptian Slit-faced Bat 

Approximate distribution of species: recorded from the Island of Corfu (Greece) 
and Palestine; Arabia; Egypt, Sudan, Kenya, Angola. 

Nycteris thebaica thebaica Geoffroy, 1818 

1818. Nycteris thebaicus E. Geoffroy, Description de I'Egypte, 2: 119, pi. i. No. 2. 

1840. Nycteris albiventer Wagner, Schreb. Saugeth. Suppl. /.• 439. Nubia, Sudan. 

Recorded from Palestine as a valid race by Aharoni, 1944, Bull. Zool. Soc. 

Egypt, 6: 26. 
Range: Egypt, Palestine, Corfu, Northern Arabia. 

Nycteris thebaica adana Andersen, 191 2 

1912. Petalia thebaica adana Andersen, Ann. Mag. N.H. 10: 548. Myba, near Aden, 
Southern Arabia. 

Genus: Megaderma, page 108 



Genus MEGADERMA E. Geoflroy, 1810 

18111. Mesiaderma GcoflVoN', Ann. Mus. H.N. Paris, /j: 190. I'espniilin spasma 

1847. Eucheira Hodgson, J. Asiat. Soc. Bengal, 16: B91. Megadcrma schistacea Hodgson 

= Megaderma Ivra Geoflroy. Not of Westwood, 1836. 
1866. Spasma Gray, P.Z.S. 83. Vespertilio spasma Linnaeus. 
1872. Lvroderma Peters, Mber. Preuss. Akad. Wiss. 195. Megaderma lyra Geoflroy. 

\"alid as a subgenus. 

2 species: Megaderma lyra, page 109 

Megaderma spasma, page 108 

We follow Chasen' and Simpson in regarding Lvroderma as of subgeneric rather 
than generic value. The two species differ in the shape of the noseleaf and also in the 
width of the skull; excellent figures are given in Dobson (1878, pi. 10). 

Subgenus MEGADERMA Geoffrey, 18 10 

Megaderma spasma Linnaeus, 1758 Malay False Vampire 

Approximate distribution of species: Ceylon, Peninsula of India, Burma, Tenas- 
serim, Cambodia (Indo-China), Siam, Malay States, Sumatra, Java, Borneo, and 
various small adjacent islands, Celebes, Philippine Islands, Ternate (Moluccas). 

(Megaderma spasma spasma Linnaeus, 1758. Extralimital) 

1758. Vespertilio spasma Linnaeus, Syst. Nat. ed. 10, /: 32. Ternate. 

Megaderma spasma horsfieldi Blyth, 1863 

1863. Megaderma horsfieldii Blyth, Cat. Mamm. Mus. Asiat. Soc. Bengal, 23. India. 
Range: Peninsula ol' India. 

Megaderma spasma medium Andersen, 1918 

1918. Megaderma spasma medium Andersen, Ann. Mag. N.H. 2: 383. Singapore 
Island. (Ranges to Tenasserim.) 

Megaderma spasm.^ majus .'\ndcrsen, 1918 

1918. Megaderma spasma majus Andersen, Ann. Mag. N.H. 2: 383. Kin, Lower 
Chindwin, Burma. 

Megaderma spasma minus Andersen, 1918 

1918. Megaderma spasma minus Andersen, Ann. Mag. X.H. 2: 383. Cambodia, Indo- 
China. Range includes Siam. 

Megaderma spasma ceylonense Andersen, 19 18 

1918. Megaderma spasma ceylonense Andersen, Ann. Mag. N.H. 2: 384. Trincomalee, 



Subgenus LYRO DERMA Peters, 1872 

Megaderma lyra Geoffroy, 18 10 Indian False Vampire 

Approximate distribution of species: Szechuan, Kwantung, Fukien, etc., in 
Southern China; India, including Bengal, Palanpur, Sikkim, Bhutan Duars, several 
localities in the Peninsula, south at least to Mysore and Western Ghats CBlanford 
gave Kashmir to Cape Comorin and Ceylon, west to Karachi^i ; Shan States, Burma; 
Malay States. 

Megaderma lyra lyra Geoffroy, 18 10 

1810. Megaderma lyra E. Geoffroy, Ann. Mus. H.X. Paris, /j.- 190. India. 1? East 

coast, Madras.) 
1839. Vesperlilio (Megaderma) carnatica Elliot, Madras J. Lit. 10: 96. Dharwar, 

Southern Mahratta, India. 
1844. Megaderma spectrum \Vagner, in Hiigels Kashmir, 569, pi. Kashmir. 
1847. Megaderma schistacea Hodgson, J. Asiat. Soc. Bengal, 16: 889. Xorth-Eastern 

Bengal, India. 

Range: Burma, Bhutan Duars, Sikkim, Bengal, Kumaon, Palanpur, Khandesh, 
Central Provinces, Bellarv', Mysore (India). 

Megaderma lyra sinensis Andersen & \Vroughton, 1907 

1907. Eucheira sinensis Andersen & ^V^oughton, Ann. Mag. N.H. jg: 136. Amoy, 

Fukien, China. 
1930. Megaderma spasma Shih, Bull. Biol. Dept. Sun. Yat-sen Univ. 9, i. Not of 

Linnaeus, 1758. (South-\Vestern border of Hunan, China.) 
Range: Chinese range of species as above, and Malay States. 

Megaderma lyra c.aurin.a Andersen & ^\'roughton, 1907 

1907. Eucheira lyra caurina Andersen & Wroughton, Ann. Mag. X.H. ig: 136. Surat 
district, India. Range includes Dharwar, Kanara and Western Ghats, 
Peninsular India. 


Genera: Asellia, page 130 

Aselliscus, page 130 
Coelops, page 131 
Hipposideros, page 123 
Rhinolophus, page 1 1 1 
Triaenops, page 131 

Of these genera, all hut Rhinolophus belong to the subfamily Hipposiderinae, which 
Miller, 1907, Families a? Genera of Bats, made a distinct family. The two groups are 
closely allied and frequently referred, as here, to a single family. 


Subfamily Rhinolophinae 

Genus RHINOLOPHUS Lacepcdc, 1799 

1790. Rhinolopkus Lacepedc, Tabl. Mamm. 15. Vespertilio ferrum-equinum Schreber. 
1836. Rkinocrepis Gervais, Diet. Pittoresque H.X. 4, 2: 617. Vespertilio ferrum-eqmnum 

1847. Aquias Gray, P.Z.S. 15. Rhinolopkus luctiis Temminck and Rhinolopkus trifoliatus 


1866. Phrllotis Gray, P.Z.S. 81. Not of W'aterhouse, 1857. Rkliiolnpkus philippinensis 

Watcrliouse. • 

1867. Coelopkvllus Peters, P.Z.S. 1866': 427. Rkinolophus cnelophvllus Peters. 

1 90 1. Euryalus Matschie, S.B. Ges. Naturf. Fr. Berlin, 225. Rhinolopkus mehelyi 

1934. Rhinophvllotis Iredale & Troughton, Mem. Austral. Mus. 6: 92. Rhinolopkus 

meoapkyllus Gray, from .Australia. (Nom. nud.) 

The most recent reviews of part of this very large genus are Tate, 1939, Amer. Mus. 
Nov. No. 1036, and 1943, Amer. Mus. Nov. No. 1219. These papers deal with the 
Oriental members of the genus, and slightly modify the arrangements of Andersen, 
1905, Ann. Mag. N.H. 16: 243, 281, 289 and 648; 1905, P.Z-S- 2: 75, 121 ; and 1918, 
Ann. Mag. N.H. 2: 374. Andersen recognized six groups of species m Rkinolophus, one 
of which appears to be extralimital, and one of which, the macrotis group, Tate 
apparently merges with the Indus . group. We entirely agree with Tate that the 
''simplex'' group of Andersen flater called "'megaphyllus" group) must be called the 
ferrumequinum group ; the last is the type species and much the earliest name in the genus. 

In the present region, the following 21 species seem most likely to prove valid: 
Rkinolophus acrotis, page 1 13 Rkinolopkus macrotis, page 122 

Rhinolopkus affinis, page 1 1 3 Rhinolopkus malayanus, page 1 1 5 

Rkinolopkus blasii, page 120 Rkinolopkus mekelyi, page 120 

Rkinolopkus clivosus, page 1 12 Rkinolophus monoceros, page 1 19 

Rkinolopkus coelophvllus, page 123 Rkinolophus pearsoni, page 122 

Rkinolopkus cornutus, page 1 1 7 Rkinolophus rex, page 1 23 

Rhinolopkus euryale, page 119 Rhinolopkus rouxi, page 114 

Rhinolopkus ferrumequinum, page 11 i Rkinolophus subhadius, page 119 
Rkinolopkus hipposideros, page i 15 Rkinolophus tkomasi, page i 14 

Rhinolopkus lepidus, page 1 1 8 Rkinolopkus trifoliatus, page 1 2 1 

Rkinolopkus luctus, page 1 2 1 

Rhinolopkus ferrumequinum group 

Tate (1939) lists four subgroups which come into the region now under discussion, 
typified by ferrumequinum, affinis, rouxi and borneensis (Rkinolophus horneensis Peters, 
1 86 1, Mber. Preuss. Akad. Wiss. 709, Labuan, North Borneo). In the present region, 
of the species listed above only R. malayanus belongs to the borneensis subgroup; 
Osgood recorded this species from Indo-China. The two principally Ethiopian species, 
R. clivosus and R. acrotis, are nearest ferrumequinum, and /?. tkomasi is near rou.xi. 


Rliinolophus ferrumequinum Schreber, 1774 Greater Horseshoe Bat 

Approximate distribution of species: England, France, Spain and Portugal, Italy, 
Switzerland, Holland, Germany, Hungary, Greece, Corsica and Sardinia; Crimea, 
Caucasus, Russian Turkestan; Japan, Korea, China (states of Chihli, Shantung, 
Shensi, Szechuan, Yunnan, Fukien) ; Asia Minor, Persia, Syria, Palestine; Kashmir, 
Kumaon, Nepal, Sikkim; Algeria, Morocco. 

Rhinolophus FERRUMEquiNUM FERRUMEQUINUM Schreber, 1774 

1774. Vespertilio ferrum-equinum Schreber, Saugeth, /; pi. 62, upper figs, (text, p. 174). 

1776. Vespertilio equinus Miiller, Natursyst. Suppl. Regist. Band, 20. France. 

1777. Vespertilio solea Zimmermann, Spec. Zool. Geogr. Quad, 452. Not available, 

see Bull. Zool. Nomencl. 4, 1950: 547. 
1779. Vespertilio perspicillatus Blumenhach, Handb. Naturgesch. 75 (part). 
1785. Vespertilio ungula Boddaert, Blench. .\nim. /.• 71. Burgundy, France. 
1792. Vespertilio ferrum-equinum major Kerr, Anim. Kingd. 99. Not of Kerr, loc. cit. 97. 

1798. Vespertilio hippocrepis Schrank, Fauna Boica, /.• 64. Renaming oi ferrum-equinum. 
1813. Rhinolophus unihastatus Geoffrey, Ann. Mus. H.N. Paris, 20: 257. France. 
(?) 1829. Rhinolophus unifer Kaup, Skizz. Europ. Thierw. /.■ 104, nom. nud. 
1863. Rhinolophus ferrum-equinum var. germanicus Koch, Jb. Nassau Ver. Naturk. 18: 

522. Wiesbaden, Germany. 
1863. Rhinolophus ferrum-equinum var. italicus Koch, loc. cit. 523. Italy. 
1885. Rhinolophus unihastatus \-a.T. homorodalmasiensisDada.Y, Orv. Term. Ert. Kolosvar, 

10: 274. Homorod-Almas Caves, Hungary. 
1887. Rhinolophus unihastatus var. homodorensis Daday, Ert. Term. Korebol, Budapest, 

16, 7: 13. Renaming oi homorodalmasiensis. 

1904. Rhinolophus ferrum-equinum obscurus Cabrera, Mem. Soc. Esp. H.N. 2: 2^J. 

Valencia, Spain. 

1905. Rhinolophus ferrum-equinum typicus Andersen, P.Z.S. igo^, 2: 113. 

191 1. Rhinolophus ferrum-equinum colchicus Satunin, Izv. Kauk. Otd. Russ. Geog. Obsc. 
21: 47-48. [N.V.) Abkhazia (Southern Russia). (Satunin, 1914, Mitt. 
Kaukas. Mus. 8: 89.) 

Range: Continental Europe, as listed above, eastwards to Russia; Algeria. (The 
form obscurus is recognized as valid by Andersen and by G. Allen (1939), from 
Spain, Balearic Islands, Algeria, Morocco.) 

Rhinolophus ferrumequinum Nippon Temminck, 1835 

1835. Rhinolophus nippon Temminck, Mon. Mamm. 2: 30a. Japan. Range includes 
Fukien, Shantung, Szechuan, etc., in China; Hokkaido, Hondo, Shikoku 
Kiushiu, Tsushima, ? Riukiu Islands, Japan. 

Rhinolophus ferrumequinum tragatus Hodgson, 1835 

1835. Rhinolophus tragatus Hodgson, J. Asiat. Soc. Bengal, 4: 699. Nepal. 
1863. Rhinolophus brevitarsus Blyth, Cat. Mamm. Mus. Asiat. Soc. Bengal, 24, nom. nud. 
Range includes Sikkim; and Yunnan, China. 


Rhinolophus FERRUMEquiNUM PROxiMus Andersen, 1905 

1905. Rhinolnphus ferrum-eqiiinum proximiis Andersen, P.Z.S. ic/Oj, 2: 112. Gilgit, 


1905. Rhinolophus Jtrriim-cquinum regains Andersen, P.Z.S. igo^, 2; 112. Mussoorie, 
Kumaon, Northern India. 

Rm.xoLOPHUs FERRUMEQUINUM iNSUL.\Nus Banett-Hainilton, 19 10 
1910. Rhinolophus ferrum-equinum insulanus Barrett-Hamilton, Ann. Mag. N.H. j.- 292. 
Cheddar, Somersetshire, England. 

Rhinolophus ferrumequinum irani Cheesman, 1921 

1 92 1. Rhinolophus ferrum-equinum irani Cheesman, J. Bombay N.H. Soc. 2y: 35. 
Shiraz, 5,200 ft., Persia. 

Rhinolophus ferrumequinum mikadoi Ognev, 1927 

1927. Rhinolophus ferrum-equinum mikadoi Ognev, J. Mamm. 8: 142. Yokohama, 
Hondo, Japan. 

Rhinolophus ferrumequinum quelpartis Mori, 1933 

1933. Rhinolophus quelpartis Mori, J. Chosen N.H. Soc. 16: i, 4. Ki-nei, Quelpart 
Island, off Korea. 

Rhinolophus ferrumequinum korai Kuroda, 1938 

1938. Rhinolophus ferrumequinum korai Kuroda, List Jap. Mamm. f|i (in full, 92). 

Southern Korea. 
1 93 1. Rhinolophus nippon pachyodonlus Kishida 6c Mori, Zool. Mag. Tokyo, 43, 379, 

nom. nud. Korea. 

Rhinolophus bocharicus Kastschenko & Akimov, 1917 

iqi7. Rhinolophus hoeharicus Kastschenko & Akimov, Annu. Mus. Zool. Acad. St. 
Petcrsb. 22: 221. Murghab River, South Russian Turkestan. Considered a 
subspecies o[ R. ferrumequinum by Ognev, 1928, Mamm. of E. Europe, N. 
Asia, /.• 397; but Kuzyakin, in Bobrinskii (1944), lists it as a full species, 
from South-Eastcrn Turkmenia, districts of Tashkent, Samarkand, near 
Kokand, and district of Termez, migrating to Afghanistan in the winter. 

Rhinolophus clivosus Crctzschmar, 1B28 

Appro.ximatr distribution of species: Red Sea coasts of Arabia and African coast 
of Gulf of Aden. 

Rhinolophi^s CLivost s Crctzschmar, 1828 

1828. Rhinolophus elirosus Crctzschmar, in Ruppcll, Atlas Reise Nordl. Afrika, 

Saut;cth. 47. Mohila, Red Sea coast, approximately 27°49' N., 35°3o' E., 



Rhinolophus acrotis Heuglin, 1861 

Approximate distribution of species: Egypt, Southern Arabia, Eritrea, the Sahara 
(in part). (B.M. specimens of this species from Hadramaut, Southern Arabia, and 
from Yemen, South-\V'estern Arabia.) 

(Rhinolophus acrotis acrotis Heuglin, 1861. Extrahmital) 

1861. Rhinolophus acrotis Heuglin, Nova Acta Leop. Carol. 2g, 8: 4, 10. Keren, 

Rhinolophus .acrotis andersoni Thomas, 1904 

1904. Rhinolophus andersoni Thomas, Ann. Mag. N.H. 14: 156. Eastern Desert of 

Egypt, about 22° N., 35° E. 

Rhinolophus acrotis brachygnathus Andersen, 1905 

1905. Rhinolophus acrotis brachygnathus Andersen, Ann. Mag. N.H. /j; 73. Giza, 


Rhinolophus acrotis schwarzi Heim de Balsac, 1934 

1934. Rhinolophus acrotis schwar~i Heim de Balsac, Bull Mus. H.N. Paris, y: 483. 
Djanet, Tassah des Azdjers, about 24°4o' N., 9°25' E., Algerian Sahara. 

Rhinolophus afiinis Horsfield, 1823 

Approximate distribution of species: Southern China (Szechuan, Yunnan, Fukien, 
Chekiang, etc.), Hainan; Kumaon, Nepal, Bhutan Duars, Darjeeling, Burma (from 
Pegu to Chindwin, at least); Tonkin, Indo-China; Malay States, Sumatra, Java, 
Natuna and Anamba Islands. 

(Rhinolophus afflnis affinis Horsfield, 1823. Extralimital) 

1823. Rhinolophus affinis Horsfield, Zool. Res. Java (6), pi. figs, a, b. Java. 

Rhinolophus affinis him.'^layanus Andersen, 1905 

1905. Rhinolophus affinis himalayanus Andersen, P.Z.S. igo^, 2: 103. Mussoorie, 

Kumaon, North-Western India. Ranges eastwards to Burma (part) and 

China (Hunan, Szechuan, Yunnan). 

Rhinolophus affinis macrurus Andersen, 1905 

1905. Rhinolophus affinis macrurus Andersen, P.Z.S. /5105, 2: 103. Taho, Karennee, 

South-Eastern Burma. Range includes Fukien and Chekiang, Southern 

China and Tonkin. 

Rhinolophus affinis tener Andersen, 1905 

1905. Rhinolophus affinis tener Andersen, P.Z.S. igo^, 2: 103. Pegu, Burma. 

Rhinolophus affinis hainanus J. Allen, 1906 

1906. Rhinolophus hainanus ]. Allen, Bull. Amer. Mus. N.H. 22: 482. Pouten, Island 

of Hainan. 



Rhinolophus andamanensis Dobson, 1872 

1872. Rhinolophus andamanensis Dobson, J. Asiat. See. Bengal, 41, 2: 337. South 

Andaman Islands, Bay of Bengal. This is very like R. affinis and may be a 

representative of it. 

Rhinolophus rouxi Temminck, 1835 

Approximate distribution of species: Ceylon, Peninsula of India, Nepal, Dar- 
jeeling, China (states of Szechuan, Yunnan, Fukien, Chekiang). 

Rhinolophus rouxi rouxi Temminck, 1835 

1835. Rhinolophus rouxii Temminck, Mon. Mamm. :?.• 30b. Pondicherry and Calcutta, 

1850. Rhinolophus rubidus Kelaart, J. Ceylon Br. Asiat. See. 2: 209. Kaduganava, 


1 85 1. Rhinolophus fulvidus Blytii (error for rubidus Kelaart), J. Asiat. Soc. Bengal, 20: 


1852. Rhinolophus cinerascens Kelaart, Prodr. Faunae Zeyl. 13. Fort Frederick, Ceylon. 
1852. Rhinolophus rammanika Kelaart, loc. cit. 14. Amanapoora Hill, Kaduganava, 

Range: Ceylon, Nilgiri Hills, Dharwar, Kanara, Nepal, Darjeeling, etc. 

Rhinolophus rouxi sinicus Andersen, 1905 

1905. Rhinolophus rouxi sinicus Andersen, P.Z.S. 2: 98. Chinteh, Anhwei, Southern 
China. Range: Chinese range of the species. 

Rhinolophus thomasi Andersen, 1905 

Approximate distribution of species: Burma, Yunnan, Tonkin. 

Rhinolophus thomasi thomasi Andersen, 1905 

1905. Rhinolophus thomasi Andersen, P.Z.S. iQOfj, 2: 100. Karin Hills, South-Eastern 

Rhinolophus thomasi latifolius Sanborn, 1939 

1939. Rhinolophus thomasi latifolius Sanborn, Field Mus. Publ. Zool. 24: 39. Muong 
Moun, Tonkin, Indo-China. 

Rhinolophus thomasi septentrionalis Sanborn, 1939 

1939. Rhinolophus thomasi septentrionalis Sanborn, Field Mus. Publ. Zool. 24: 40. 
Nguluko, 27 '5' N., ioo"i5' E., north of Likiang, Yunnan, China. 

Tate lists the fillDwing little-known species in the rouxii subgroup. 

Rhinolophus petersi Dobson, 1872 

1872. Rhinolophus prtcrsii Dobson, J. Asiat. Soc. Bengal, 41, 2: 337. No locality. 

Perhaps Irum India. Blanford, 1891, listed it from Mussoorie, and Coonoor 

in the Nilgiii Hills. 



Rhinolophus malayanus Bonhote, 1903 

Approximate distribution of species: Lower Siam and Indo-China (Tonkin). 

Rhinolophus malayanus Bonhote, 1903 

1903. Rhinolophus malayanus Bonhote, Fasc. Malayenses, Zool. /; 15. Biserat, Jalor, 

Malay Peninsula. 

Other named form: 

Rhinolophus chaseni Sanborn, 1939 

1939. Rhinolophus chaseni Sanborn, Field Mus. Publ. Zool. 24: 38. Pulau Condor 

(Condor Island), off Southern Indo-China. From description, nearest 


Rhinolophus hipposideros group 

Andersen originally called this the "midas group", but subsequently adopted the 
above name (correctly so, since hipposideros antedates by more than a hundred years). 

Rhinolophus hipposideros Bechstein, 1800 Lesser Horseshoe Bat 

Appro.ximate distribution of species: England, Ireland, France, Spain, Portugal, 
Switzerland, Italy, Sardinia, Corsica, Malta, Germany, Poland, Hungary, Russia 
(Southern Ukraine, Caucasus); South Russian Turkestan; Asia Minor, Persia, 
Cyprus, Arabia; Kashmir; Morocco; Sudan, Eritrea. 

Rhinolophus hipposideros hipposideros Bechstein, 1800 

1792. Vespertilio ferrum-equinum minor Kerr, Anim. Kingd. 99, not minor Kerr, loc. cit. 

97. France. 
1800. Vespertilio hipposideros Bechstein, in Pennant, Uebers. Vierf Thiere, 2: 629. 

1813. Rhinolophus bihastatus Geoffroy, Ann. Mus. H.N. Paris, 20: 259. Neighbour- 
hood of Paris, France. 
(?) 18 1 6. Phyllorhina minuta Leach, Syst. Cat. Mamm. & Birds B.M. 5, nom. nud. 
(?) 1829 Rhinolophus bifer Kaup, Skizz. Europ. Thierw. /.• 104, nom. nud. 
1840. Rhinolophus bifer B\^L\nviUe , Osteographie, Vespertilio, 31. 
1863. Rhinolophus hipposideros var. typus Koch, Jb. Nassau Ver. Naturk, 18: 530. 

Wiesbaden, Germany. 
1863. Rhinolophus hipposideros var. alpinus Koch, loc. cit. Alps. 
1870. Rhinolophus eggenhbffner Fitzinger, S.B. Akad. Wiss. Wien, 61, i: 151. MS. 

synonym of bihastatus. 
1885. Rhinolophus bihastatus var. kisnyiresiensis Daday, Orv. Term. Ert. Kolozsvar, 10: 

274. Kis-Nyires, Szolnok Dobaka, Hungary. 
1887. Rhinolophus hipposideros var. trogophilus Daday, Ert. Term. Korebol, Budapest, 

16, 7: 8. Renaming oi kisnyiresiensis. 

1904. Rhinolophus euryale helvetica Bretscher, Vischr. Naturf Ges. Zurich, 4g: 256. 

Baar, Zug, Switzerland. 

1905. Rhinolophus hipposideros tjpicus Andersen, P.Z.S. igo§, 2: 141. 




(?) 1920. Rhinolophus anomalus Sodcrlund, Zool. Anz. 5i\- 122. Wildbad Gastcin, 

Salzburg, Austria. 
(?) 1920. Rhinolophiis intnrnedius Sodcrlund, loc. at. 124. Wildbad Gastein, Salzburg, 

(?) 1943. Rhinolophus moravicus Kostron, Acta Soc. Sci. Nat. Moravia, Brno, 75, 9: 13. 

Moravia, Czechoslovakia. See also Kostron, 1946, Casopis Vlast. Spolkn. 

Mus. Olmutz, f)^: i-ii. 
(?) 1943. Rhinolophus hipponderoi intermedius Laurent, Bull. Soc. Z. France, 68: 188. 

Not of Sodcrlund, 1920. Geneva, Switzerland. 
Range: Continental Europe, north of the Alps, through Armenia to North-Western 

Rhinolophus hipposideros minutus Montagu, 1808 

1808. Vespcrtilio minutus Montagu, Trans. Linn. Soc. London, g: 163. Wiltshire, Ranges to Ireland. 

Rhinolophus hipposideros minimus Heuglin, 1861 

1861. Rhinolophus minimus Heuglin, Nova Acta Leop. Carol. 2g, 8: 6. Keren, 

Eritrea, North-Eastcrn Africa. 
1863. Rhinolophus hipposideros var. pallidus Koch, Jb. Nassau \'er. Naturk. 18: 531. 

Mediterranean region. 

1904. Rhinolophus phasma Cabrera, Mem. Soc. Esp. H.N. 2: 252. Madrid, Spain. 
Range: Mediterranean region (quoted by Miller from Spain, Portugal, France, 

Switzerland, Italy, Corsica, Sardinia, Malta, Cyprus); also Eritrea and Senaar, 
Sudan (G. Allen); recorded from Arabia (Taif) by Morrison-Scott (1939). 

Rhinolophus hipposideros midas Andersen, 1905 

1905. Rhinolophus wiWaj Andersen, /905, 2: 138. Jask, Persian Gulf Range: Gilgit to 

Cyprus, according to Andersen (1918), who appears to treat this form as a 
subspecies in his key (p. 378) where its status, and that of the other named 
forms recognized, seems not very clear. 

Rhinolophus hipposideros m.«iJORi Andersen, 1918 

1918. Rhinolophus hipposideros majori Andersen, Ann. Mag. N.H. 2: -^11, 378. Patri- 
nionio. Northern Corsica. 

Rhinolophus hipposideros escaler,^e Andersen, 19 18 

1918. Rhinolophus hipposideros escalerae Andersen, .'\nn. Mag. N.H. 2: 378. Ha-ha, 
Mogador, Morocco. 

Rhinolophus hipposideros vespa Laurent, 1937 

1937. Rhinolophus hipposideros vespa Laurent, Bull. Soc. H N. Afr. N. 28: 157 Korifla, 

Rhinolophus pusillus group 
Andersen first called this the "lepidus group" (1905), subsef|uently the pusillus 
group. Tate prefers the first, and lists one of its subgroups as the ''minor subgroup". 



But minor Horsfield, 1823, from Java, is preoccupied by minor Kerr, 1792 = hippo- 
sideros, and so cannot be used in this group. R. pusillus Temminck, 1834, is the next 
available name for minor Horsfield [nee Kerr) and appears to be the earliest name in 
the group. The type locality for pusillus is Java, and we believe this species to be 
wholly extralimital to our list, notwithstanding the fact that under the name "minor" 
it was listed by earlier authors from Darjeeling and Siam. The few skins examined 
from Java are all unusually dark in colour and easily distinguished from such species 
as cornutus or blythi, which represent the group on the mainland, and the latter of 
which is likely to occur in Darjeeling and Siam. But we suggest that there is very little 
evidence that blythi is in reality a species distinct from cornutus, as we suspect the 
dental details given by Andersen to separate blythi may not be constant, and there is 
no difference in size (as judged by forearm length) between the two supposed species 
when all races are taken into account. 

Tate divided the Oriental members of this group into three subgroups, typified by 
pusillus {"minor"), lepidus and subbadius, and in addition to these, the three well-known 
European species, blasii, euryale and mehelyi belong here. These have been compared 
with the Oriental species by Andersen. R. monoceros belongs to the subbadius sub- 

The reference oi R. pusillus is Temminck, 1834, Tijdschr. Nat. Gesch. Phys. i: 29 

Rlunolophus cornutus Temminck, 1835 Little Japanese Horseshoe Bat 

Approximate distribution of species : Japan ; Liukiu Islands; Szechuan, Fukien, 
Kwantung, Hainan, etc., in China; Indo-China; Siam; Kumaon, India; and 

Rhinolophus cornutus cornutus Temminck, 1835 

1835. Rhinolophus cornutus Temminck, Mon. Mamm. 2: 37. Japan. Range includes 
Hokkaido, Hondo, Shikoku, Kiushiu, Iki Islands, Tsushima. 

Rhinolophus cornutus pumilus Andersen, 1905 

1905. Rhinolophus cornutus pumilus Andersen, P.Z.S. igo§, 2: 127. Okinawa, Liukiu 
Islands. Range includes Szechuan and Kwantung, China. 

Rhinolophus cornutus perditus Andersen, 19 18 

1918. Rhinolophus perditus Andersen, Ann. Mag. N.H. 2: 376. Ishigaki, Southern 
Liukiu Islands. 

Rhinolophus cornutus blythi Andersen, 1918 

1918. Rhinolophus blythi Andersen, Ann. Mag. N.H. 2: 376, 377. Almora, 5,500 ft., 
Kumaon, Northern India. 

Rhinolophus cornutus szechwanus Andersen, 1918 

1 9 18. Rhinolophus blythi szechwanus Andersen, Ann. Mag. N.H. 2: 376, 377. Chunking, 

Szechuan, China. Range: Szechuan, Hupeh, Yunnan, Burma, Darjeeling, 




Rhinolophus cornutus calidus G. Allen, 1923 

1923. Rhinolophus blj'thi calidus G. Allen, Amer. Mus. Nov. No. 85 ; i. Yenping, Fukien, 

South-Eastern China. Ranges to Tonkin, Indo-China. 

Rhinolophus cornutus orii Kuroda, 1924 

1924. Rhinolophus cornutus orii Kuroda, New Manim. Riukiu Islands, 4. San-Mura, 

Tokunoshima, 300 ft., Liukiu Islands. 

Rhinolophus cornutus miyakonis Kuroda, 1924 

1924. Rhinolophus miyakonis Kuroda, New. Mamm. Riukiu Islands, 5. Nishisato, 
Miyakojima, Liukiu Islands. 

Rhinolophus cornutus parous G. Allen, 1928 

1928. Rhinolophus blythi parcus G. Allen, Amer. Mus. Nov. No. 317: 2. Nodoa, Island 
of Hainan. 

Andersen regards the following member of the pusillus subgroup as a distinct 
species : 

Rhinolophus gracilis Andersen, 1905 

1905. Rhinolophus gracilis Andersen, P.Z.S. 2: 129. Malabar coast, India. 

Rhinolophus lepidus Blyth, 1844 

Approximate distribution of species: Szechuan and Yunnan, China; Central 
Provinces, Ganges Valley, Kumaon, Bengal, etc., in India; Mt. Popa, Pagan and 
Chindwin River, Burma. 

Rhinolophus lepidus lepidus Blyth, 1844 

1844. Rhinolophus lepidus Blyth, J. Asiat, Soc. Bengal, ij: 486. ? Calcutta. Range: 
India, as above. 

Rhinolophus lepidus shortridgei Andersen, igi8 

1918. Rhinolophus lepidus shortridgei Andersen, Ann. Mag. N.H. 2: 376, 377. Pagan, 

Burma. Range includes Chindwin, Burma; also Szechuan and Yunnan, 


The following species, probably belonging to the lepidus subgroup, have also been 

Rhinolophus monticola Andersen, 1905 

1905. Rhinolophus monticola Andersen, P.Z.S. igo§, 2: 124. Mussoorie, Kumaon, 
North-Western India. 

Rhinolophus feae Andersen, 1907 

1907. Rhinolophus feae Andersen, Ann. Mus. Stor. Nat. Genova, j: 474. Biapo, Karin 
Hills, Burma. 


Rhinolophus OSGOOD! Sanbom, 1939 

1939. Rhinolophus osgoodi Sanborn, Field Mus. Publ. Zool. 24: 40. Nguluko, 27°5' N., 
ioo°i5' E., north of Likiang, Yunnan, China. 

Tate Hsts the following in the lepidus subgroup, but according to Andersen's key 
(1918) they belong to the garoensis (= subbadius) subgroup. 

Rhinolophus cognatus cognatus Andersen, 1906 

1906. Rhinolophus cognatus Andersen, Ann. Mus. Stor. Nat. Genova, 3, 2: 181. Port 
Blair, South Andaman Islands, Bay of Bengal. 

Rhinolophus (?) cognatus famulus Andersen, 19 18 

1918. Rhinolophus famulus Andersen, Ann. Mag. N.H. 2: 377. North Central Island, 
Andaman Islands, Bay of Bengal. 

Rhinolophus subbadius Blyth, 1844 

Approximate distribution of species: Nepal, United Provinces (India) and Assam; 
Tonkin, Indo-China. 

Rhinolophus subbadius Blyth, 1B44 

1841. Rhinolophus subbadius Hodgson, J. Asiat. Soc. Bengal, 10: 908, nom. nud. 

1844. Rhinolophus subbadius Blyth, J. Asiat. Soc. Bengal, 13: 486. Nepal. 

1872. Rhinolophus garoensis Dobson, J. Asiat. Soc. Bengal, 41, 2: 337. Garo Hills, 

Assam. Andersen ( 1918) lists garoensis as a valid form, but does not compare 

it with subbadius. Wroughton listed it as a synonym. 

Rhinolophus monoceros Andersen, 1905 
Distribution: Formosa. 

Rhinolophus monoceros Andersen, .1905 

1905. Rhinolophus monoceros AndeTstn, P.Z.S. igo^, 2: 131. Baksa, Formosa. 

Rhinolophus euryale Blasius, 1853 Mediterranean Horseshoe Bat 

Approximate distribution of species: Portugal, Spain, France, Italy, Sardinia, 
Austria, Yugoslavia, Greece; south-east coast of Black Sea, Caucasus, and South- 
West Russian Turkestan (Turkmenia) ; Syria, Palestine (Asia Minor, according to 
Kuzyakin) ; Morocco, Algeria, Egypt. 

Rhinolophus euryale euryale Blasius, 1853 

1853. Rhinolophus euryale Blasius, Arch. Naturgesch. ig, 1: 49. Milan, Italy. 

1904. Euryalus toscanus Andersen & Matschie, S.B. Ges. Naturf. Fr. Berlin, 77. 

Caverna di Parignana, Mt. Pisani, Italy. 
1904. Euryalus atlanticus Andersen & Matschie, loc. cit. St. Paterne, Indre-et-Loire, 

1904. Euryalus cabrerae Andersen & Matschie, loc. cit. 78. Alcala de Henares, Madrid, 




Rhixolophi's euryale barbarus Andersen & Matschic, 1904 

ig04. Einvalus barbarus Andersen & Matschie, S.B. Ges. Naturf. Fr. Berlin, 79. 

Tans;iers, Morocco. 
I?) 1867. Rhirwlophus algiriis Loche, Expl. Sci. dc TAl^crie, Zool. Mamm. 8;^,. Altreria. 

Ranges eastwards to Tunis. 

Rhinolophus euryai.e meridionalts Andersen & Matschie, 1904 
1904. Eurvalus mcridtonalis Andersen & Matschic, S.B. Ges. Xaturf. Fr. Berhn, 70. 
Algeria ("probably a mountain form"). 

Rhinolophus eury.\le jud.mcus Andersen & Matschic, 1904 
1904. Eiirvalus jiidaiais Andenen & Matschie, S.B. Ges. Naturf. Fr. Berlin, 80. Cave 
of Adullam, Jerusalem, Palestine. Range: to Egypt. 

Rhinolophus euryale nordmanni Satunin, 191 1 

191 1. Rhinolophus eurvale nordmanni 'S)3.t\in\n, Izv. Kavkaz. Otd. R.G.O. 21: 47. (N.V.) 
Pa\lovsk, Sukhum district, Transcaucasia. 

Rhinolophus mehelyi Matschie, 1901 

Approximate distribution of species: Spain, Southern France, Sardinia, Rumania, 

Rhinolophus mehelyi Matschie, 1901 

1901. Rhinolophus mehelvi Matschie, S.B. Ges. Naturf Fr. Berlin, 225. Bucharest, 

1904. Rhinolophus carpelanus Cabrera, Mem. Soc. Esp. ?i.N. 2: 254. Madrid, Spain. 

Rhinolophus blasii Peters, 1866 

Approximate distribution of species: Greece, Cyprus, Italy (whence recorded in 
1931); Palestine; Transcaucasia and Turkmenia (South-\\'est Russian Turkestan); 
Asia Minor (according to Kuzyakin) ; North Africa (Dobson) ; and in G. Allen's 
Checklist African Mamm., but without details. 

Rhinolophus blasii Peters, 1866 

1857. Rhinolophus divosus Blasius, Siiugeth. Deutschlands, 33. Not of Cretzschmar, 

1828. (Italy, Sicily, Istria, Dalmatia.) 
1866. Rhinolophus blasii Peters, Mber. Preuss. Akad. Wiss. 17. New name for divosus 

Blasius nee Cretzschmar. 
1910. Rhinolophus hlasiusi Trouessart, Faunc Mamm. dT.urope, 9. 

Rhinolophus luelus group 

Andersen (1911-)) originally called this the philippinensis group (based on R. philippi- 
nensis \Vaterhouse, 1843, P-Z-^- 68, from Luzon), but later ( 1918) he renamed it the 
luetus group. Strictly, it should be known as the trifoliatus group, as Irifoliatus ante- 
dates luelus bv one vear. However, in order not to introduce further nomenciatural 


muddle, we retain the name luctus for the group. Tate, 1943, Amer. Mus. Nov. 
No. 1 2 19, has considerably altered Andersen's arrangement of this group. Tate 
divides the group into three sections, typified by luctus, trifoliatus and philippinensis; to 
the section typified by the latter he apparently refers macrotis, coelophjUus and rex. 

Rhinolophus trifoliatus Temminck, 1834 Trefoil Horseshoe Bat 

Approximate distribution of species : Darjeeling, Tenasserim, South-Western Siam, 
Malay States, Sumatra, Java, Borneo, and adjacent small islands. 

Rhinolophus trifoliatus trifoliatus Temminck, 1834 

1834. Rhinolophus trifoliatus Temminck, Tijdschr. Natuur. Gesch. /.• 24, pi. i, fig. 6. 

The following very little known form is listed near trifoliatus by Tate, but 
Wroughton regarded it as unidentifiable. 

Rhinolophus mitratus Blyth, 1844 

1844. Rhinolophus mitratus Blyth, J. Asiat. Soc. Bengal, 75.- 483. Chaibassa, Orissa, 

Rliinolophus luctus Temminck, 1835 Great Eastern Horseshoe Bat 

Approximate distribution of species: Tenasserim, Burma, Nepal, Sikkim, United 
Provinces, Peninsular India, Ceylon; Fukien (in South-Eastern China), Hainan, and 
probably represented Formosa; Malay States, Sumatra, Java, Borneo. 

Tate (1943) appears to regard all named forms as subspecies, but we have retained 
R. pearsoni as distinct because it seems to occur with luctus, and it differs from it in size. 

Rhinolophus luctus luctus Temminck, 1835 

1835. Rhinolophus luctus Temminck, Mon. Mamm. 2: 24, pi. 30. Java. Occurs to 
Tenasserim, according to Wroughton; this might be the form Rhinolophus 
morio Gray, 1842, Ann. Mag. N.H. 10: 257, from Singapore, a valid race 
according to Chasen (1940). 

Rhinolophus luctus perniger Hodgson, 1843 

1843. Rhinolophus perniger Hodgson, J. Asiat. Soc. Bengal, 12: 414. Nepal. Range 
includes Kumaon, Sikkim; Chin Hills and Shan States, Burma. 

Rhinolophus luctus lanosus Andersen, 1905 

1905. Rhinolophus lanosus Andersen, Ann. Mag. N.H. 16: 248. Kuatun, North- 
W'estern Fukien, China. 

Rhinolophus luctus beddomei Andersen, 1905 

1905. Rhinolophus beddomei Andersen, Ann. Mag. N.H. 16: 253. Wynaad, Madras, 
India. Range; Peninsula of India. 



Rhinolophus luctos sobrinus Andersen, 1918 

1918. Rhinolophus heddomei sobrinus Andersen, Ann. Mag. N.H. 2: 378. Kala Oya, 
North Central Pro\'incc, Ceylon. 

Rhinolophus luctus spurcus G. Allen, 1928 

1928. Rhinolophus lanosus spurcus G. Allen, Amer. Mus. Nov. No. 317: 3. Nodoa, 
Island of Hainan. 

Rhinolophus (?) luctus formosae Sanborn, 1939 

1939. Rhinolophus formosae Sanborn, Field Mus. Publ. Zool. 24: 41. Formosa. 

Rhinolophus pearsoni Horsfield, 1851 

Approximate distribution of species: Kumaon, Darjeeling, Assam (Dobson); 
Szechuan, Yunnan, Fukien, in China; Indo-Clhina (Tonkin). 

Rhixolophus pearsoni pearsoni Horsfield, 1851 

1851. Rhinolophus pearsoni Horsfield, Cat. Mamm. Mus. E. Ind. Co. 33. Darjeeling, 

North-Eastern India. 
1872. Rhinolophus larvatus Milne-Edwards, Rech. H.N. Mamm. 248, pi. 37a, fig. i; 

pi. 37c, fig. I. Not of Horsfield, 1823. Moupin, Szechuan, China. 
1872. Rhinolophus yunanensis Dobson, J. Asiat. Soc. Bengal, 41, 2: 336. Hotha, 

Yunnan, China. 
Range : as above, except Tonkin and Fukien. 

Rhinolophus pearsoni chinensis Andersen, 1905 

1905. Rhinolophus pearsoni chinensis Andersen, Ann. Mag. N.H. 16: 289. Kuatun, 
Fukien, South-Eastern China. Range: to Tonkin. 

Rhinolophus macrotis Blyth, 1844 Large-eared Horseshoe Bat 

Approximate distribution of species: Szechuan and Fukien, China; Kumaon, 
Nepal; Indo-China; Sumatra; Philippine Islands (Tate). 

Rhinolophus macrotis macrotis Blyth, 1844 

1844. Rhinolophus macrotis Biyth, J. Asiat. Soc. Bengal, i^: 485. Nepal. 

Rhinolophus m.\crotis siamensis Gyldenstolpe, 1916 

1916. Rhinolophus macrotis siamensis Gyldenstolpe, K. Svenska Vetensk. Akad. Handl. 
^j, 2: 12. Doi Par Sakeng, North-Western Siam. Range: to Tonkin, Indo- 

RiiiNOLOPHrs macrotis episcopus G. Allen, 1923 

1923. Rhinolophus episcopus G. Allen, Amer. Mus. Nov. No. 85: 2. VVanhsicn, Szec- 
huan, China. (Tate (1943) makes this a race oi macrotis.) 

Rhinolophus macrotis caldwelli G. Allen, 1923 

1923. Rhinolophus episcopus caldwelli G. Allen, Amer. Mus. Nov. No. 85: 3. Yuki, 
Fukien, China. Range: to Tonkin, Indo-China. 



Rhinolophus coelophyllus Peters, 1867 

Approximate distribution of species: Burma, Siam, Malay States. 

Rhinolophus coelophyllus coelophyllus Peters, 1867 

1867. Rhinolophus coelophyllus Peters, P.Z.S. 1866: 426, pi. 35. Salween River, Burma. 

(Known from Moulmein and Tsagine in Upper Burma, Malay States, and 

Chiengmai, Siam (Tate).) 

Rhinolophus coelophyllus shameli Tate, 1943 

1943. Rhinolophus coelophyllus shameli Tate, Amer. Mus. Nov. No. 1219: 3. Koh Chang 
(Island), Siam. 

Rhinolophus rex G. Allen, 1923 

.\ppro.\imate distribution of species: Szechuan and Kweichow, China. 

Rhinolophus rex G. Allen, 1923 

1923. Rhinolophus rex G. Allen, Amer. Mus. Nov. No. 85: 3. Wanhsien, Szechuan, 

Subfamily Hipposiderinae 

Genus mPPOSIDEROS Gray, 1831 

183 1. Hipposideros Gray, Zool. Misc. 37. Vespertilio speoris Schneider. 

1837. Phyllorhina Bonaparte, Fauna Ital., pt. 21 : 3. Rhinolophus diadema E. Geoffroy. 

1866. Gloionycteris Gray, P.Z.S. 82. Phyllorhina armiger Hodgson. 

1866. Speorifera Gray, P.Z.S. 82. Hipposideros vulgaris Blyth —Rhinolophus laruatus 

1866. Chrysonycteris Gray, P.Z.S. 82. Hipposideros fuhms Gray. 

1866. Rhinophylla Gray, P.Z.S. 82. Phyllorhina labuanensis Tomes. Not of Peters, 1865. 

1866. Macronycteris Gray, P.Z.S. 82. Rhinolophus gigas Wagner, from Angola. 

1 87 1. Doryrhina Peters, Mber. Preuss. Akad. Wiss. 314. Phyllorhina cyclops Temminck, 
from the Gold Coast. 

1 87 1. Sideroderma Peters, Mber. Preuss. Akad. Wiss. 324. Phyllorhina fuliginosa Tem- 
minck, from \Vest Africa. 

1871. Ptychorhina Peters, Mber. Preuss. Akad. \V'iss. 325. Rhinolophus caffer Sundevall. 

1871. Cyclorhina Peters, Mber. Preuss. Akad. Wiss. 326. Phyllorhina obscura Peters, 
from Luzon, and P. doriae Peters, from Borneo. 

1 87 1. Thyreorhina Peters, Mber. Preuss. Akad. Wiss. 327. Phyllorhina coronata Peters, 
from Mindanao, Philippine Islands. 

1871. Synodesmotis Peters, Mber. Preuss. Akad. Wiss. 329. Phyllorhina megalotis Heug- 
lin, from Eritrea. 

1888. Hipposiderus Blanford, P.Z.S. i88j: 637 (Emendation). 

This genus is revised in some detail by Tate, 1941, Bull. Amer. Mus. N.H. j8: 
353-393, who divides the genus into 1 1 species groups, six of which occur in the 
present region. 

I 123 

PALAEARC;TIC and IXDIAN mammals 1758-1946 

In the present region, the following 1 1 species seem most likely to prove valid: 

Hippoiideros armigcr, page 128 
Hipposideros bicolor, page 126 
Hipposideros coffer, page 129 
Hipposideros cineraceus, page 127 
Hipposideros diadema, page 125 
Hijiposideros galcritus, page 1 29 
Hipposideros lankadiva, page 125 
Hipposideros larvatus, page 124 
Hipposideros pomona, page 127 
Hipposideros pratti, page 129 
Hipposideros speoris, page 1 24 

Hipposideros speoris group 

H. speoris is the earliest name in the genus. Tate (1941, 377, 378) compares the two 
species referred here. 

Hipposideros speoris Schneider, 1800 Schneider's Leaf-nosed Bat 

,\pprnxiniatc distribution of species: Ceylon, Peninsula of India. Has also been 
recorded (possibly erroneously) from Java, Borneo, Timor. 

Hipposideros speoris speoris Schneider, 1800 

1800. Vespertilio speoris Schneider, in Schreber's Saugeth., pi. 59b. Tranquebar, 

India. (Tate, 1941, Bull. Amer. Mus. N.H. y8: 377.) 
1 83 1. Rhinolophus diikhunensis Sykes, P.Z.S. 99. Deccan, India. 
1838. Hip/iiisideros apiculatus Gray, Mag. Zool. Bot. 2: 492. ^[adras, India. 
1838. Hipposideros penicillatus Gray, loc. cit. 493. Madras, India. 
1850. Hipposideros tcmpletonii Kelaart, J. Ceylon Br. Asiat. Soc. 2: 208. Ceylon. 
i8'^2. Hipposideros aureus Kelaart, Prodr. Faun. Zeylan, 18. Ceylon. 
1852. Hipposideros hlvthi Kelaart, loc. cit. 20. 
Range: Ceylon; Dharwar, Kanara, Mysore, Coorg, etc., in Peninsular India. 

Hipposideros speoris puixhellus Andersen, 1918 

19 1 8. Hipposideros speoris pulcliellus Andersen, .\nn. Mag. N.H. 2: 383. Vijayanagar, 
Bellary, India. 

Hipposideros larvatus Horsfield, 1823 

.VpprDximate distribution of species: Hainan; Assam, Burma; Indo-China; Malay 
States, Sumatra, Java, Borneo. 

Hipposideros l.^rvatus larvatus Hursfield, 1823 

1823. Rhinolophus larvatus Horsfield, Zool. Res. Java, No. 6, pi. 9. Java. Recorded 

from Tonkin, Indo-China, by Osgood; Chasen, however, seems to restrict 

this form to Java. 




1874. Phyllorhina leptophylla Dobson, J. Asiat. Soc. Bengal, ^j, 2: 234. Khasi Hills, 


1906. Hipposideros poutensis ]. Allen, Bull. Amer. Mus. N. H. 22: 483. Pouten, Island 
of Hainan. 

HiPPOSIDEROS LARVATUS gr.'>lNdis G. Allen, 1936 

1936. Hipposideros larvatus grandis G. Allen, Rec. Ind. Mus. ^5, 3: 345. Akanti, Upper 
Chindwin, 500 ft., Burma. ("Not improbably a synonym of leptophylla" 



1942. Hipposideros larvatus alongensis Bourret, C. R. Conseil Rcch. Sci. Indochine, 
ig42, 2: 27. Bay d'Along, Indo-China. 

Hipposideros diadema group 

The subgeneric name Phyllorhina is available here, if subgeneric division is required. 
The two well-known species referred here are discussed by Tate (1941); see also 
Andersen, 19 18, Ann. Mag. N.H. 2: 381. 

Hipposideros diadema E. Geoffroy, 1813 Large Malay Leaf-nosed Bat 

Approximate distribution of species : Burma, Indo-China, Malay States, Java, 
Sumatra, Borneo, and some adjacent small islands, Celebes, Philippine Islands; 
perhaps represented in New Guinea, Queensland, Solomon Islands, etc. 

(HiPPOSIDEROS DIADEMA DIADEMA E. Geoffroy, 1813. Extralimital) 

1813. Rhinolophus diadema Geoffroy, Ann. Mus. H.N. Paris, 20: 263, pi. 6. Island of 
Timor. Range includes Java. 

Hipposideros diadema masoni Dobson, 1872 

1872. Phyllorhina masoni Dobson, J. Asiat. Soc. Bengal, 41, 2: 338. Moulmein, Burma. 
Range includes Annam, Indo-China. 

Hipposideros lankadiva Kelaart, 1850 

Approximate distribution of species: Ceylon, Peninsula of India. 

Hipposideros lankadiva lankadiv.\ Kelaart, 1850 

1850. Hipposideros lankadiva Kelaart, J. Ceylon Br. Asiat. Soc. 2: 216. Kandy, Ceylon. 

Hipposideros lankadiva indus Andersen, 1918 

1918. Hipposideros indus Andersen, Ann. Mag. N.H. 2: 382. Gersoppa, Kanara, 
Peninsular India. 



HiPPOSiDEROS LANKADivA MixTUs Andersen, 1918 

1918. Hipposideros Indus mixtus Andersen, Ann. Mag. N.H. 2: 382. Kolar, Eastern 
Mysore, India. 

Hipposideros lan'Kadiva u.mtus Andersen, 191 8 

1918. Hipposideros indus unitus Andersen, Ann. Mag. N.H. 2: 382. Mundra, Sanger, 
Central Provinces, 1,600 ft., India. 

Other named species in the diadema group (it is possible that nicobarensis represents 
diadema and that schistaceus represents lankadiva) : 

Hipposideros nicobarensis Dobson, 1871 

1871. Phrllorhina nicobarensis Dobson, J. Asiat. Soc. Bengal, ^o, 2: 262. Nicobar 
Islands, Bay of Bengal. 

Hipposideros schistaceus Andersen, 1918 

1918. Hipposideros schistaceus Andersen, Ann. Mag. N.H. 2: 382. Vijayanagar, 
Bcllary, India. 

Hipposideros bicolor group 

Revision: Andersen, 1918, Ann. Mag. N.H. 2: 379. Several species are admitted, 
all of which seem closely allied to each other. H. cineraceus, the smallest in size, 
appears valid. Tate (1941, 363) lists gentilis and allied forms as races of bicolor, but 
Chasen, 1940, Bull. Raffles Mus. i§: 44, regards bicolor and gentilis as species occur- 
ring together. We suggest pomona is the earliest name for the races currently referred 
to gentilis. There seems little evidence that the Indian fulvus is in reality more than 
western subspecies of bicolor. 

Chrrsonrcteris Gray is available if subgcncric division is required. 

Hipposideros bicolor Temminck, 1834 Bicoloured Leaf-nosed Bat 

.\pproximatc distribution of species: Nicobar Islands, Condor Island (off Cochin- 
China) ; Lower Siam, Sumatra, Java; as here understood, also Ceylon, Peninsula of 
India ("where widely distributed), Sind, Cutch, Rajputana, Kathiawar; Sikkim, 
Bhutan Duars, Burma fChindwin to Shan States, Mt. Popa), Tenasserim; Formosa 


Hipposideros bicolor bicolor Temminck, 1834 

1834. Rhinolophus bicolor Temminck, Tijdschr Natuur. Gesch^ /, i: 19, pi. i,_fig. 3. 

.^njer coast. North- Western Java (Tate) 
Siam, Sumatra, Java. 

Range: Condor Island, Lower 



HipposiDEROs (?) BicoLOR FULVus Gray, 1838 

1838. Hipposideros fulvus Gray, Mag. Zool. Bot. 2: 492. Dharwar, India. 

1838. Hipposideros murinus Gray, Mag. Zool. Bot. 2: 492. Madras, India. 

1839. Rhinolophus fulgens Elliot, Madras J. Lit. 10: 99. Dharwar, India. 
1859. Phyllorhina aurita Tomes, P.Z.S. 76. India. 

Range: Peninsula of India, as far north as Nasik, Bombay. 

Hipposideros (?) bicolor ater Templeton, 1848 

1848. Hipposideros ater Templeton, J. Asiat. Soc. Bengal, 77, i: 252. Colombo, 

1850. Hipposideros atratus Kelaart, J. Ceylon Br. Asiat. Soc. 2: 208. Colombo, Ceylon. 

Substitute for ater. 

Hipposideros bicolor nicobarulae Miller, 1902 

1902. Hipposideros nicobarulae Miller, Proc. U.S. Nat. Mus. 24: 781. Little Nicobar 
Island, Bay of Bengal. 

Hipposideros (?) bicolor pallidus Andersen, 1918 

1 91 8. Hipposideros fulvus pallidus Andersen, Ann. Mag. N.H. 2: 381. Junagadh, 
Kathiawar, India. Range: Kathiawar, Cutch, Sind, Rajputana, India. 

Hipposideros pomona Andersen, 1918 

Approximate distribution of species : Coorg, India, and if gentilis is correctly 
allocated here, Burma; Fukien, Yunnan, in China; Hainan, Indo-China; Siam, 
Malay States, islands west of Sumatra (Nias and Engano), Java, Banka. 

Hipposideros pomona pomona Andersen, 1918 

1 918. Hipposideros pomona Andersen, Ann. Mag. N.H. 2: 380, 381. Haleri, North 
Coorg, Southern India. 

Hipposideros pomona gentilis Andersen, 191 8 

1 918. Hipposideros gentilis Andersen, Ann. Mag. N.H. 2: 380, 381. Thayetmyo, 
Burma. Ranges to Tonkin and Annam, in Indo-China. 

Hipposideros pomona sinensis Andersen, 191 8 

1918. Hipposideros gentilis sinensis Andersen, Ann. Mag. N.H. 2: 380, 381. Foochow, 
Fukien, Southern China. Range includes Yunnan and Hainan. 

Hipposideros cineraceus Blyth, 1853 

Approximate distribution of species: Punjab, India; Burma; Raheng, in Siam; 
Tonkin, in Indo-China; Malay States, Rhio Archipelago, Borneo, Anamba Islands. 

Hipposideros cineraceus cinEraceus Blyth, 1853 

1853. Hipposideros cineraceus Blyth, J. Asiat. Soc. Bengal, 22: 410. Near Pind Dadan 
Khan, Salt Range, Punjab. Range: as above. 



1872. PhvUorhina micropus Peters, Mber. Preuss. Akad. Wiss. 256. Dehra Dun, near 
Simla, Xorth-\\'cstern India. 

The species //. amhoinensis Peters, 1871, Mber. Preuss. Akad. Wiss. 323, from 
Amboina Island (Moluccas), which Tate says is probably a synonym of aruensis 
Gray, i8-,8, P.^S. 107, Aru Islands, off New Guinea, was recorded from parts of 
India by earlier authors: Blanford (1891), Dobson (1878) and Wroughton (1918). It 
is unlikely that an Australasian bat would occur in islands off New Guinea, India, 
and nowhere eke. Dobson placed micropus in the synonymy oC amhoinensis, and it is 
most likely that ■'amhoinensis" of the earlier writers on Indian Chiroptera is the 
species now called cineraceus. 

Hipposideros armiger group 
The subgencric name Gloionvcteris is available for this group. 

Hipposideros armiger Hodgson, 1835 Great Himalayan Leaf-nosed Bat 

Approximate distribution of species: Szechuan, \"unnan, Fukien and adjacent 
states in .South-Eastern C:hina; Formosa, Liukiu Islands; Kumaon, Nepal, Assam, 
Burma iChin Hills, Shan States, Mt. Popa, etc.); Tonkin, in Indo-China; Malay 

Hipposideros armiger ar.miger Hodgson, 1835 

1835. Rhmolophus armiger Hodgson, J. Asiat. Soc. Bengal, 4: 699. Nepal. Ranges from 
Kumaon to Burma, Tonkin, Yunnan and Szechuan, China. 

Hipposideros armiger swinhoei Peters, 1871 

1871. Phrllorhina swmhoii Peters, in Swinhoe, P.Z.S. i8jo: 616. Amoy, Fukien, China. 
Ranges to Kiangsu and Chekiang, South-Eastern China. ("Seems to be in- 
distinguishable Irom armiger" (Tate, 1941, 390).) 

Hipposideros (?) armiger turpis Bangs, 1901 

K)Oi. Hipposideros turpu Bangs, Amer. Nat. 35: 561. Ishigaki, South Liukiu Islands. 

Hipposideros armiger debilis Andersen, 190b 

ir)o6. Hipposideros armiger dehilis Andersen, .^nn. Mag. N.H. ij: 37. Province 

Welleslev, Malay Peninsula. Perhaps extra limital to this list, but according 

to Tate reaches Siam. 

Hipposideros armiger terasensis Kishida, 1924 

1924. Hipposideros armiger terasensis Kishida, Zool. Mag. Tokyo, j6: 42. Formosa. 
l.^'.r.) "Seems to be indistinguishable from armiger" (Tate, 1941, 390). 

Hipposideros armiger tranninhensis Bourret, 1942 

1942. Hipposideros tranninhensis Bourret, C.R. Conseil Rcch. Sci. Indochine, ig42, 2: 
20. Jarres, Tran-Ninh, Indo-China. 



Hipposideros galeritus group 

Tate refers H. coffer, from Africa, to the present group, and for this the name 
Ptychorhina is available if subgeneric division is required. 

Hipposideros galeritus Cantor, 1846 

Approximate distribution of species: Ceylon, Bengal, Southern Bombay, Palanpur, 
Central India; ? Assam, ? Burma; Malay States, Sumatra, Borneo, and certain small 
adjacent islands. 

Hipposideros galeritus galeritus Cantor, 1846 

1846. Hipposideros galeritus Cantor, J. Asiat. Soc. Bengal, 75.- 183. Penang, Malay 
States. Tate, 1947, Mamm. E. Asia, quotes it from Burma and Assam. 

Hipposideros galeritus brachyotus Dobson, 1874 

1874. Phyllorhina brachyola Dobson, J. Asiat. Soc. Bengal, ./j, 2: 237. Central India. 

Range: Ceylon, Bengal, Kanara, Palanpur, Central India. Tate (1941, 367) 

suggests it is a race oi galeritus. 

Hipposideros caffer Sundevall, 1846 South African Lesser Leaf-nosed Bat 

Approximate distribution of species: Morocco, and south of the Sahara, from 
Eritrea and Kenya, and from Gabon district, at least, southwards to South-West 
Africa, Natal, the Transvaal, and Pondoland in Eastern Cape Province. South- 
western Arabia, vide Hayman, 1941, in Brit. Mus. Exp. S.W. Arabia, 1937-8, 
Chiroptera, 2. 

(Hipposideros caffer caffer Sundevall, 1846. Extralimital) 
1846. Rhinolophus caffer Sundevall, Ofvers. Vetensk. Akad. Forh. Stockholm, j, 4: 
118. Near Durban, Natal, South Africa. 

Hipposideros caffer tephrus Cabrera, 1906 

1906. Hipposideros tephrus Cabrera, Bol. Soc. Esp. H.N. 6: 358. Mogador, Morocco. 

For notes on the characters of the caffer subgroup, see Tate (1941, 366). 
Hipposideros pratti group 

Hipposideros pratti Thomas, 1891 Pratt's Leaf-nosed Bat 

Approximate distribution of species: China, states of Szechuan, Fukien, Chekiang; 
Shan States, in Burma; Siam; Tonkin, in Indo-China; Malay States. 

Hipposideros pratti pratti Thomas, 1891 

1891. Hipposiderus (sic) pratti Thomas, Ann. Mag. N.H. j: 527. Kiatingfu, Szechuan, 
China. Range: China, as above, and Tonkin. 



1913. Hipposideros Ijlei Thomas, Ann. Mag. N.H. 12: 88. Chiengdao Cave, 50 
miles north of Chlengmai, Northern Siam. Range: Burma, Siam, Malay 

Genus ASELLIA Gray, 1838 

1838. Ast-llia Gray, Mag. Zool. Bot. 2: 493. Rhinolophus tridens Geoffroy. 

The subsidiary genera of Hipposiderinae, Asellia, Aselliscus, Triaenops, Coelops, and 
a few others, were reviewed by Tate, 1941, Amer. Mus. Nov. No. 1140. Asellia is 
restricted by Tate to A. tridens only. 

I species: Asellia tridens, page 130 

Asellia tridens E. Geoffroy, 1813 Trident Leaf-nosed Bat 

Approximate distribution of species: Sind, India; Arabia, Iraq, Palestine; Egypt, 
Algeria, Morocco; southwards in Africa to Somaliland and Zanzibar. Blanford also 
quoted it from Southern Persia. 

Asellia tridens tridens E. Geoffroy, 1813 

1 81 3. Rhinolophus tridens Geoffroy, Ann. Mus. H.N. Paris, 20: 265. Egypt. Range: 
Egypt to Zanzibar (G. Allen). 

Asellia tridens murraiana J. Anderson, 1881 

1881. Phyllorhina tridens var. murraiana Anderson, Cat. Mamm. Ind. Mus. 113. 
Karachi, Sind, Western India. 


1918. Asellia tridens diluta Andersen, Ann. Mag. N.H. 2: •^'ji^. El Golea, Algerian 
Sahara. Range includes Biskra, Northern Algeria. 

Asellia tridf.ns pallida Laurent, 1937 

1937. Asellia tridens pallida Laurent, Mammalia, /.• iii. Oued Tatta, Anti-Atlas, 
South-Western Morocco. 

Genus ASELLISCUS Tate, 1941 

1941. Aselliscus Tate, Amer. Mus. Nov. No. 1140: 2. Rhinolophus tncuspidatus Tcm- 
minck, from Amboina (Moluccas). 

\Vhen Tate erected this genus he suggested that the species stolic^kanus (Dobson, 
1 87 1, from Penang) and trifidus should be referred to it. He also compared with them 
the species ^'Triaenops'' or '^Asellia" wheeleri, and came to the conclusion that the latter 
was closely related to stoltczkanus, and that in skull characters wheeleri represents a 
"quite advanced Hipposiderine near Aselliscus'\ As wheeleri seems distinct from both 


Asellia and Triaenops, it is tentatively referred here. See also Dorst, 1948, Mammalia, 
12: 16. We do not know whether wheeled is a valid species or a race of one of the 
earlier-named species just quoted. 

? 2 species in the area covered by this list: 

Aselliscus trifidus, page 131 
Aselliscus wheeleri, page 131 

Aselliscus trifidus Peters, 1871 

Approximate distribution of species: Burma. 

Aselliscus trifidus Peters, 1871 

1 87 1. Phyllorhina trifida Peters, P.Z.S. 513. Burma. 

Aselliscus wheeleri Osgood, 1932 

Approximate distribution of species: Indo-China; Kweichow, in Southern China; 
Northern Burma. 

Aselliscus wheeleri Osgood, 1932 

1932. Triaenops wheeleri Osgood, Field Mus. Publ. Zool. 18: 224. Muong Moun, 
Tonkin, Indo-China. 

Genus TRIAENOPS Dobson, 1871 
1871. Triaenops Dobson, J. Asiat. Soc. Bengal, 40, 2: 455. Triaenops persicus Dobson. 
I species in the area covered by this list: 
Triaenops persicus, page 131 

Triaenops persicus Dobson, 1871 Persian Leaf-nosed Bat 

Approximate distribution of species: Persia, Arabia, south to Aden, Egypt. 

Triaenops persicus Dobson, 1871 

1 87 1. Triaenops persicus Dobson, J. Asiat. Soc. Bengal, 40, 2: 455, pi. 18. Shiraz, 
about 4,750 ft., Persia. Range: to Aden and Egypt. 

Genus COELOPS Blyth, 1848 

1848. Coelops Blyth, J. Asiat. Soc. Bengal, ly, i: 251. Coelops frithii Blyth. 
191 1. Chilophylla Miller, Proc. U.S. Nat. Mus. 38: 395. Chilophylla hirsuta Miller, 
from Mindoro, Philippine Islands. 

I species in the area covered by this list: 

Coelops frithi, page 132 

From descriptions it seems that there is not likely to be more than one species in 
the area now under consideration. 



Coelops frithi BIyth, 1848 Tailless Leaf-nosed Bat 

Approximate distribution oi' species: Szcchuan and Fukien, China; Formosa; 
Bengal; Tonkin and Annani, Indo-China; Java, and perhaps Malay Peninsula (if 
rohinsorii Bonhote, 1908, is a race o^ frithi). 

Coelops frithi frithi Blyth, 1848 

1848. Coelopi frithii Blyth, J. Asiat. Soc. Bengal, ij: 251. Sundarbans, Bengal, India. 

Coelops frithi inflatus Miller, 1928 

1928. Coelops injlata Miller, Proc. Biol. Soc. \Vashington, 41: 85. Ycnpingfu, 2,000 ft. 
Fukien, South-Eastern China. Range: to Indo-China. 

CoELOP.s frithi sinicus G. Allen, 1928 

1928. Coelops sinicus G. Allen, Amer. Mus. Nov. No. 317: 4. Two miles north-east of 
Wanhsicn, Szechuan, C'hina. 

Coelops frithi for.mos.'vnus Horikawa, 1928 

1928. Coelops formosnrms Horikawa, Trans. N.H. Soc. Formosa, j8, No. 98: 339. 
Kuraru, in Koshun, Formosa. 


Genera: Ulnmops, page 136 
Til da I i da, page 1 32 

Genus TAD ARID A Rafincsque, 181 4 

1 8 14. Tadarida Rafincsque, Precis Som. 55. Cephalotes Icniolis Rafincsque. 

1818. Js'yclinomus E. Geoflroy, Description de I'Egypte, i\- 1 14. Nyctiiwmus aenyptiacus 

1 82 1. Nvctinoma Bowdich, Anal. Nat. Class. Mamm. 28. 

1 82 1. .Nyctinomes Gray, London Med. Repos. 75.- 299. 

1822. Nyctinomia Fleming, Philos. Zool. 2: 178. 

1825. Dinops Savi, N. Giorn. Lett. Pisa, Sci. 10: 229. Dinops cestoni Sa\i = Cephalotes 

temotis Rafincsque. 
1830 vcl 1 83 1. Dvsnpes Cretzschmar, in Rtippcll, Atlas Rcisc nordl. Afrika, Saugeth. 

69. Not of Illiger, 1811. 
1842. Mops Lesson, Nouv. Tabl. Rcgn. Anim. 18. \[ops indiais Lesson = Dysopes 

mops F. Cuvier, from Sumatra. Valid as a subgenus 
1865. Mormopterus Peters, Mber. Preuss. Akad. Wiss. 258. Nyctinomus jugidaris Peters, 

from Madagascar = Vespertilio acetabulosus Hermann from Mauritius. Valid 

as a subgenus. 
1874. Chaerephon Dobson, J. Asiat. Soc. Bengal, ^j, 2: 144. Nyctinomus johorensis Dob- 
son, from johore, Malay States. Valid as a subgenus. 
1902. Nyctinomops Miller, Proc. Acad. Nat. Sci. Philadelphia, 393. Nyctinomus 

Jemorosaeca Merriam, from California. 



1917. Lophomops ]. Allen, Bull. Amer. Mus. N.H. ^j: 460. Chaerephon {Lophomops) 

chapmi ]. Allen, from the Belgian Congo. 
191 7. AllomopyJ. Allen, Bull. Amer. Mus. N.H. jj: 470. Chaerephon (Allomops) 

osborni J. Allen, from the Belgian Congo. 
1934. Philippinopterus Taylor, Philippine Land Mamm. 314. Philippinopterus lanei 

Taylor, from the Philippine Islands. 
1934. Micronomus Iredale & Troughton, Mem. Austral. Mus. 6: 100. Molossus 

norfolcensis Gray, from Norfolk Island (Australasia). (Nom. nud.) 
1934. Austronomus Iredale & Troughton, loc. cit. Molossus australis Gray, from New 

South Wales, Australia. (Nom. nud.) 

This genus was formerly called Njctinomus by virtually all zoologists, but Tadarida 
antedates. Thomas & Hinton, 1923, P-Z-S- 251, would separate Nyciinomus (type 
aegyptiacus) from Tadarida (type teniotis) on account of the presence of four or six lower 
incisors respectively. Miller, however, did not consider this of even subgeneric value. 
It is customary to divide this genus, which has a nearly world-wide range, into half a 
dozen or more "genera". Tate, 1941, Amer. Mus. Nov. No. 1142, has shown that 
the chief character used by Miller, 1907, Families & Genera of Bats, 244, in his key to 
the genera, to divide the genera into groups is not strictly constant in Chaerephon. 
Thomas, 19 13, J. Bombay N.H. Sac. 22: 89-91, rearranged the genera and compli- 
cated the classification by yet further generic splitting. We cannot help feeling that 
Simpson (1945) is correct in stating that the groups Chaerephon, Mops and Mormopterus, 
which we have included above as of subgeneric value, can well be included in the 
genus Tadarida. Simpson also included Otomops in the genus Tadarida, but we adopt 
Mr. R. W. Hayman's suggestion {in lilt.) that Otomops should be retained as a full 
genus on account of its remarkable cranial characters and striking external features. 

5 species of Tadarida in the area covered by this list: 

Tadarida aegvptiaca, page 134 
Tadarida plicata, page 135 
Tadarida pumila, page 1 35 
Tadarida teniotis, page 133 
Tadarida tragata, page 135 

For key to species, see Dobson, 1878, Cat. Chiroptera, 420. 

Subgenus TADARIDA Rafinesque, 1814 

Tadarida teniotis Rafinesque, 1814 European Free-tailed Bat 

Approximate distribution of species: Portugal, France, Italy, Sicily, Greece — has 
been recorded from Switzerland; according to Kuzyakin, in the U.S.S.R. it only 
occurs in Transcaucasia and in Russian Turkestan (near Bokhara); Korea; Fukien, 
Chihli and Yunnan, China; and has been recorded from Japan and Formosa. 
Trouessart quoted it from Persia, and Bodenheimer (1935) from Palestine; Egypt. 
In addition, Ognev (1927) quoted it from Vladivostock. 



Tadarida teniotis teniotis Rafincsque, 1814 

1814. Cephalotes teniotis Rafinesque, Precis. Som. 12. Sicily. 

1825. Dinops cestoni Savi, N. Giorn. Lett. Pisa, Sci. 10: 235. Pisa, Italy. 
1840. Dvsopes savii Schinz, Europ. Fauna, /.• 5. Substitute for cestoni. 

1 87 1. Dinops cestonii var. nigrogriseus Schneider, N. Denkschr. Schweiz. Gcs. Naturw. 

2./, 4: 5. Basel, Switzerland. 
1 89 1. Nyctinomus taeniotis Thomas, P.Z.S. 182. 

1897. Dvsopes midas Schulze, Helios, Berlin, /./.• 95. Not of Sundevall, 1842. 
Range: Italy, Sicily, Greece, Portugal, Caucasus, Turkestan. 

Tadarida teniotis ruppelli Temminck, 1826 

1826. Dvsopes riipelii (sic) Temminck, Mon. Mamm. /.• 224, pi. 18. Egypt. 

Flower, 1932, Notes on Recent Mammals of Egypt, P.^S- 369, does not list the 
species. G. Allen ( 1939) includes it in the African list as Mopi riippelli, with a note that 
"there seems no doubt that this name must replace midas Sundevall"''! 1842, from the 
Anglo-Egyptian Sudan). There are no specimens in the British Museum, but in our 
copy of Temminck's work Thomas has noted "= teniotis". Allen does not include 
Tadarida teniotis in his African list. Mr. R. W. Hayman, who has compared the skulls 
of teniotis and midas with Temminck's description and figures of riippelli, informs us 
that there is no doubt that Thomas was right and that Allen was wrong in listing 
midas, which is a true Mops, as a synonym q( riippelli. 


1861. Nvctinomus insignis Blyth, J. Asiat. Soc. Bengal, 50.- 90. Amoy, Fukien, China. 
1870. Dvsopes (Molossus) rueppelii Swinhoe, P.Z.S. 619. Not of Temminck, 1826. 
1920. Tadarida latouchei Thomas, Ann. Mag. N.H. fj: 283. Chiiigwantao, coast of 

North-Eastern Chihli, China. 
1 93 1. Tadarida septentrionalis Kishida, in Kishida & Mori, Z. Mag. Tokyo, ^5.' 379, 

nom. nud. (J^.V.). N. Korea. 
Range: Fukien and Chihli, in China; Korea and Ussuri region; Japan (Abe, 1944). 

Tadarida teniotis coecata Thomas, 1922 

1922. Tadarida teniotis coecata Thomas, Ann. Mag. N.H. 10: 392. Mekong Valley, 
about 28^20' N., 7,000 ft., Yunnan, China. 

Tadarida aegyptiaca E. Geoffroy, 1818 

Appro.ximate distribution of species: Egypt; Kenya (Hollister, 1918) ; Sind, Cutch, 
Poona, Rajpiitana, P.ilanpur, Kalliiawar, Mysore, Dharwar and Deccan, India; 
Znluland and Cape Province (Roberts). 
Tadarida aegyptiaca aegyptiaca E. Geoffroy, 1818 

1 818. Nvctinomus aegvptiacus Geoffroy, Description de I'Egypte, 2: 128, pi. 2, No. 2. 

1826. Dvsopes geoffrovi Temminck, Mon. Mamm. /.• 226, pi. 19. Substitute for 




The following also appear to be subspecies : 

Tadarida aegyptiaca sindica Wroughton, 191 9 

1919. Tadarida sindica Wroughton, J. Bombay. N.H. Soc. 26: 732. Kashmor, Upper 
Sind Frontier, India. 

Tadarida aegyptiaca thomasi Wroughton, 1919 

1919. Tadarida thomasi Wroughton, J. Bombay N.H. Soc. 26: 732. Bhuj, Cutch, 

Tadarida aegyptiaca gossei Wroughton, 19 19 

1919. Tadarida gossei Wroughton, J. Bombay N.H. Soc. 27; 733. Sassoon Hospital, 
Poona, India. 

Tadarida tragata Dobson, 1874 

Appro-ximate distribution of species: Calcutta and Malabar, India. (Wroughton 
(1919) stated that all but one specimen in the B.M. from India for this subgenus 
belong to the aegyptiaca section.) 

Tadarida tragata Dobson, 1874 

1874. J^yctinomus tragatus Dobson, J. Asiat. Soc. Bengal, 43, 2: 143. Calcutta, India. 

Subgenus CHAEREPHON Dobson, 1874 

Tadarida plicata Buchannan, 1800 Wrinkle-lipped Bat 

Approximate distribution of species: Rajputana, Peninsula of India, Ceylon, 
Tenasserim; Hainan; Malay States, Sumatra, Borneo, Java; probably represented 
in the Philippine Islands and Northern Australia. 

Tadarida plicata plicata Buchannan, 1800 

1800. Vespertilio plicatus Buchannan, Trans. Linn. Soc. London, 5: 261, pi. 13. 

Bengal, India. 
1820. Nyctinomiis bengal ensis DesTmLTest, Encyclop. Meth. (Mamm.), /.• 116. 
1830. Dysopes murinus Gray, Illustr. Ind. ZooL, pt. 3, pi. i. 

Tadarida plicata insularis Phillips, 1932 

1932. Chaerephon plicatus insularis Phillips, Spolia Zeylan. 16: 334. Kumbalgamuwa, 

3,000 ft., near Mulhalkelle, 30 miles south-east of Kandy, Central Province, 


Tadarida pumila Cretzschmar, 1830 vel 1831 

Approximate distribution of species: three specimens in B.M. from Sabiya, 
1 7° 10' N., 42°3o' E., Arabia. South of Sahara, known from Eritrea, Southern Sudan, 
Uganda, Kenya, Angola, Portuguese East Africa, Transvaal, Bechuanaland, etc. 



Tadarida piMiLA PLMILA Cit'tzschmai", 1830 \el 1831 

1830 \cl 1831. Dvsopes fiiimihis Crctzsclimar, in Ruppcll Atlas, Rcisc Noidl. Alrika, 
Saugfth. 69, pi. 2~ . Massawa, Eliitrcii. Ranges to Araliia, as above. 

Genus OTOMOPS Thomas, 191 3 
1913. Olomops Thomas, J. Bombay X.H. See. 22: 91. Nvctinoinus wroiig/Uoni Thomas. 
I species in the area covered by this hst: 
Otomopi wrouijhtoni, page 136 

Ototnops wroughtoni Thomas, 19 13 Wroushton's Free-tailed Bat 

Approximate distribution of species: Kanara, Southern India. 

Otomops wroughtoni Thomas, 19 13 

1913. Nvctinomus wroiighloni Thomas, J. Bombay X.H. Soc. 22: 87. Barapede Cave, 
near Talewadi, Kanara, Indi.i. 

Cheiromdes (Horsfield, 1824, ^. Res. Java), with species Cheiromcles torquatus Hors- 
field, 1824, loc. cit., Penang, Malay States (the Naked Bat), was recorded from 
Indo-China by Wagner (1855) and from some part of Siam by Boitard (1842), but 
has not to our knowledge been collected in any part of the region now under discus- 
sion in recent years, and is most likely extralimital to this list. 

Distribution: Malay States, Sumatra, Java, Borneo, Celebes, Philippine 


Genera: Barhastella, page 175 .Nyctalus, page 158 

Discopus, page 151 Njcticeius, page 176 

Eptesicus, page 153 Otonyckris, page 180 

Glischropus, page 173 Pipisirellus, page 161 

Harpiocephalus, page 187 Plecotiis, page 180 

Hesperoplenus, page 173 Scotomanes, page 177 

Kcnvoula, page 187 Scotophilus, page 178 

Mimoptcrus, page 182 Tylonrcluis, page 174 

Murina, page 184 I'espertUio, page 151 
Mrolis, page 137 

This family is world-xside in distribution and one of the largest in the class 
Mammalia. Dobson (1878) gave a key to most of the species then known, but the 
nomenclature and generic arrangement of this work is now out of date. Miller, 1907, 
Families & Cenera of Bats, re\ised the genera (and oversplit them considerably); for 



key, see pp. 197-200. Simpson (1945) has attempted some reduction of Miller's long 
list of genera, but in our opinion has gone rather too far, and he lists Nyctalus Bowdich, 
1825, in Pipislrellus Kaup, 1829, although Myctalus (which is in any case a distinct 
genus) antedates by four years. Tate, 1941, Bull. Amer. Mus. N.H. j8: 567-597, has 
reviewed the Oriental members of the Miniopterinae, Kerivoulinae and Murininae, 
and 1942, Bull. Amer. Mus. N.H. 80: 221-297, the Oriental Vespertilioninae; see 
also Tate, 1941, Bull. Amer. Mus. N.H. y8: 537, on Eurasian Myoiis. On the European 
species, see Miller, 191 2, Catalogue of the Mammals of Western Europe, 165. 

Subfamily Vespertilioninae 

Genus MYOTIS Kaup, 1829 

1829. Myotis Kaup, Skizz. Europ. Thierw. /.• 106. Vespertilio myotis Borkhausen. 

1829. Nystactes Kaup, Skizz. Europ. Thierw. /.• 108. Not of Gloger, 1827. Vespertilio 

bechsteinii Kuhl. 

1830. Leuconoe Boie, Isis, Jena, 256. Vespertilio daubentonii Kuhl. Valid as a subgenus. 
1 841. Selysius Bonaparte, Faun. Ital. /.• Introd. 3. Vespertilio mystacinus Kuhl. Valid 

as a subgenus 

1 841. Capaccinius Bonaparte, loc. cit. i: Indice Distrib. i. Vespertilio capaceinii Bona- 


1842. Trilatitus Gray, Ann. Mag. N.H. 10: 258. Included three species: hasseltii 

Temminck, from Java; macellus Temminck, from Borneo; and blepotis (a 

Miniopterus) . 
1849. Tralatitus Gervais, Diet. Univ. H.N. 13: 213, modification of Trilatitus. 
1856. Brachyotus Kolenati, Allg. Dtsch. Naturh. Ztg. 2: 131. Not of Gould, 1837. 

Vespertilio mystacinus Kuhl. 
1856. Lotus Kolenati, Allg. Dtsch. Naturh. Ztg. 2: 131. Vespertilio nattereri Kuhl 

(Tate, 1 941). Valid as a subgenus. 

1866. Tralatitius Gray, Ann. Mag. N.H. ly: 90, modification oi Trilatitus. 

1867. Pternopterus Peters, Mber. Preuss. Akad. VViss. 706. Vespertilio [Pternopterus) 

lobipes Peters ? = Vespertilio muricola Gray. 
1870. Exochurus Fitzinger, S.B. Akad. Wiss. Wien, 62: 75. (Based on macrodactylus 

Temminck, horsfieldii Temminck, from Java, and macrotarsus Waterhouse, 

from the Philippine Islands.) 
1870. Aeorestes Fitzinger, S.B. Akad. Wiss. Wien, 62, i : 427. (Based on villosissimus, 

albescens Geoffroy, and nigricans \Vied, the last two from South America.) 
1870. Comastes Fitzinger, S.B. Akad. Wiss. Wien, 62, i: 565 (included Vespertilio 

capaceinii Bonaparte and Vespertilio dasycneme Boie). 
1899. Euvespertilio Acloque, Faune de France, Mamm. 38 (included emarginatus, 

murinus = myotis, mystacinus, nattereri and bechsteinii). 
1910. Chrysopteron ]enx.\uk. Notes Leyden Mus. 32.- 74. Kerivoula weberi }, from 

Celebes. Valid as a subgenus. 
191 7. Rickettia Bianchi, .'Vnnu. Klus. Zool. Acad. St. Petersb. 21: Ixxxii. Vespertilio 

(Leuconoe) ricketti Thomas. Valid as a subgenus. 
1917. Dichromyotis Bianchi, .Vnnu. Mus. Zool. Acad. St. Petersb. 21: Ixxviii. Vesper- 
tilio formosus Hodgson. 



Mvoris [conld.] 

iQiy. Paramxotis Bianchi, Amiu. Mus. Zool. Acad. St. Petcrsb. 21: Ixxix. New name 

for Nystactes Kaup, 1829, preoccupied. Vespertilio bechsteinii Kuhl. Valid as a 


It is also probable that Pizonyx Miller, 1906 (North America) and Cisltigo Thomas, 
19 1 2 (Africa) should be referred to this genus as subgenera. 

For a very able review of this genus, see Tate, 1941, Bull. Amer. Mus. N.Ii. y8: 
537. Tate recognizes seven subgenera which are adopted here. It is not quite clear 
how many species should be retained in the region now under discussion, but Tate's 
paper and other works suggest that the following 20 are most likely to be valid, in 
the area covered by this list : 

Myotis adversus, page 149 Mrotii Jrater, page 142 

Mvotis altarium, page 142 Myotis ifconnikovi, page 141 

Myotis bechsteini, page 143 Myotis macrodactylus, page 150 

Myotis blythi, page 145 Myotis myotis, page 144 

Myotis capaccinii, page 148 Myotis mystacinus, page 138 

Myotis dasycneme, page 150 Myotis nattereri, page 143 

Myotis daubentoni, page 147 Myotis pequinius, page 149 

Myotis davidi, page 149 Myotis ricketti, page 150 

Myotis emarginatus, page 141 Afyotis sicanus, page 146 

Myotis formosus, page 146 Myotis siligorensis, page 142 

We suggest that blythi is the prior name for the European oxygnathus. 

Subgenus SELYSIUS Bonaparte, 1B41 

In the present subgenus Tate recognizes three sections, typified by M. mystacinus, 
M. emarginatus and M. siligorensis, and does not allocate M. Jrater (which seems very 
distinct). There is little doubt that from descriptions M. altarium is a valid species. 
Tate listed .\I. ikonnikovi as a race of mystacinus, but it is retained following Kuzyakin, 
because it seems to occur with mystacinus in North-Eastern Asia. Where the two occur 
together, ikonnikovi averages smaller than mystacinus. 

Myotis mystacinus Kuhl, 18 19 Whiskered Bat 

Approximate distribution of species: England, Ireland, Sweden, Norway, Belgium, 
France, Spain, Switzerland, Germany, Denmark, Poland, Holland, Bohemia, Hun- 
gary, Rumania, Bulgaria ("entire Continent of Europe" according to Miller). 
Russia, north to about 62-63 'N ., south to the Black Sea and Caucasus, Russian 
Turkestan, eastwards across Siberia to the Ussuri region, Sakhalin, Kamtchatki. 
Japan, Mongolia, Chinese Turkestan, Formosa, Korea; China, states of Szechuan, 
Shansi, Chihli, Yunnan, Fukien; Persia (Ognev), Afghanistan (Kuzyakin) ; Kashmir, 
Punjab, Nepal, Sikkim, Bhutan Duars, Tenasserim; Laos, in Indo-China; repre- 
sented .Malay States, Sumatra, Java, Borneo. 

Our listing of this species is based on that of Tate, 1941- 



Myotis mystacinus mystacinus Kuhl, 1819 

1819. Vespertilio mystacinus Kuhl, Ann. Wetterau Ges. Naturk. 4, 2: 202. Germany. 

1 82 1. Vespertilio collaris Schinz, Das Thierreich von Cuvier, /.- 177. Mt. Blanc, 

Haute-Savoie, France. 
1834. Vespertilio humeralis Baillon, Mem. Soc. £mul. Abbeville, 18^;}: 50. Abbeville, 

Somme, France. 
1837. Vespertilio schinzii Brehm, Ornis, j.- 27. Renthendorf, Thuringia, Germany. 

1843. Vespertilio schrankii Wagner, Arch. Naturgesch. g, 2: 25. ? Munich, Germany. 
1863. Brachyotus mystacinus var. nigricans Koch, Jb. Nassau Ver. Naturk. 18: 444. 

Wiesbaden, Nassau, Germany. 
1863. Brachyotus mystacinus var. rufofuscus Koch, lac. cit., same locality. 
1863. Brachyotus mystacinus var. aureus Koch, loc. cit. 445. Breisgau, Germany. 
1869. Vespertilio mystacinus var. nigricans Fatio, Faune Vert. Suisse, /.■ 92. Switzerland. 
1869. Vespertilio lugubris Fatio, Faune Vert. Suisse, /.• 93. Alternative for nigricans 

1 87 1. Vespertilio mystacinus var. nigrofuscus Fitzinger, S.B. Akad. ^Viss. Wien, 65, i: 

217. Renaming oi schinzii Brehm. 
Range: Europe. 

Myotis mystacinus brandti Eversmann, 1845 

1845. Vespertilio brandtii Eversmann, Bull. Soc. Nat. Moscou, 18, i : 505. Foothills of 

Ural Mountains, U.S.S.R. 
1905. Vespertilio mystacinus sibiricus Kastschenko, Observations on mammals from 
W^. Siberia & Turkestan, in Trans. Tomsk Univ. 27, i : 25. Siberia. 

Myotis mystacinus muricola Gray, 1846 

1 841. Vespertilio muricola Hodgson, Calcutta J.N. H. 2: 212, nom. nud. 

1846. Vespertilio muricola Gray, Cat. Hodgson Coll. B.M. 4. Nepal. 

(?) 1867. Vespertilio (Pternopterus) lobipes Peters, Mber. Preuss. Akad. Wiss. 706. 

Akyab, Arakan, Burma. 
Range: Nepal to Bhutan Duars, Tenasserim and Laos. 

Myotis mystacinus caliginosus Tomes, 1859 

1859. Vespertilio caliginosus Tomes, P.Z.S. 73. India. Range: known from Simla and 

1 87 1. Vespertilio blanfordi Dobson, Proc. Asiat. Soc. Bengal, 214. Himalayas. 

Myotis mystacinus nipalensis Dobson, 1871 

1844. Vespertilio pallidiventris Hodgson, Calcutta J.N. H. 4: 286, nom. nud. 

1871. Vespertilio nipalensis Dobson, Proc. Asiat. Soc. Bengal, 214. Katmandu, Nepal. 
(?) 1926. Myotis meinertzhageniThomas, Ann. Mag. N.H. ly: 609. Junction ofNubra 

and Shyok Rivers, Ladak, Kashmir. 

Myotis mystacinus moupinensis Milne-Edwards, 1872 

1872. Vespertilio moupinensis Milne-Edwards, Rech. H.N. Mamm. 253, pi. 37a, fig. 2; 

pi. 37c, fig. 4. Moupin, Szechuan, China. Ranges to Yunnan and Fukien, 

K 139 

PAi.Ai-.ARcrnc; and indian mammals 1758-1946 

Myotis mvstacinus MONTivAGus Dobsoii, 1874 

1874. Vi-spnlilio montivagus Dobson, J. Asiat. Soc. Bcnsral, _/jj, 2 : 237. Hotlia, Yunnan, 
China. Ranajes to Chihii, Fukicn (part), China. 

Myotis mysiwcini's PRZEVVALSKn Bubrinskii, 192G 

1926. Mvotis mrslncima pizeivahkii Bobrinskii, C.R. Acad. Sci. U.R.S.S. 95. \'allcy of 

Moldja River, northern slope of Kotan Tagh, Southern Sinkiang. 
RansTC : to Shansi, China, and Russian Asia. 

Myotis mystacim's gracilis Ognev, 1927 

1927. Mrotis mnlacimis ortuilis Ognev, J. Mamm. 8: 145. Vladivostock, Eastern 

Siberia. Range includes Lake Baikal district to Sakhalin, Kamtchatka, also 
Korea, Hokkaido, Hondo, Kurile Islands. 

Myotis mystacinus transcaspicus Ogncv & Hcptncr, 1928 

1928. Mvolis mystacinus transcaspicus Ognev & Hcptncr, Zool. Anz. 75.- 260. 

Mikhailovskoi, Kopet Dag, Transcaspia. 

Myotis mystac;inu.s kukunoriensis Bobrinskii, 1929 

i()29. Mratis niYStacinus kukunoriensis Bobrinskii, Annu. Mus. Zool. Acad. St. Petcrsb. 

o^;; 221. Balekut-Gomi, Hwang Ho, south of Kukunor, Xorth-Eastcrn 


Myotis mystacinus latirostris Kishida, 1932 

1932. Myotis latirostris Kishida, Lansania, 4, 40: 153. (N.V.) Central Formosa. 
1935. Myotis muricola orii Kuroda, J. Mamm. r6: 290. Taihezan, Taihokusiu, 
Northern Formosa. 

Myotis my.stacinus sogdianus Kuzyakin, 1934 

1934. Mvolis mystacinus sogdianus Kuzyakin, Bull. Soc. Nat. Moscou, pj: 321, 329. 

Tashkent, Russian Turkestan. 

Myotis mystacinus pamirensis Kuzyakin, 1935 

1935. Myotis mvstacinus pamirensis Kuzyakin, Bull. Soc. Nat. Moscou, 44: 431, 436 

)as('hul-Kul Lake, Pamir Mountains South-East Russian Turkestan). 

Myotis .mystacinus aurascens Kuzyakin, 1935 

1935. Myotis mystacinus aurascens Kuzyakin, Bull. Soc. Nat. Moscou, 44: 432, 437. 
Knrkushin, \'ladika\kaz, Northern Caucasus. 

Myotis my.stacinus bui.garicus Hcinrich, 1936 

I(I3(). Myotis mystacinus bulgaricus Heinrich, Mitt. Naturw. List. Sofia, f): 38. East of 
Plovdiv, Bulgaria. 

Mvoiis mystacinus hajastanicus Argyropulo, njii) 

I93(). Mrotn mvstaciuNS hajastanicus Argyropulo, Zool. Pap. Piinl. Inst. Erivan, /.■ 27. 
S( hiirdsa (Nadcshino), Lake Sevanga, Armenia. 



Myotis ikonnikovi Ognev, 191 2 

Approximate distribution of species : from the Russian Altai and North-Eastern 
Mongolia, east to Sakhalin, north to Southern Yakutia, and includes Korea, 
Manchuria and Hokkaido. 

Myotis ikonnikovi Ognev, 1912 

1912. Myotis ikonnikovi Ognev, Annu. Mus. Zool. Acad. St. Petersb. 16: 477. Iman 
district, Ussuri Valley, Eastern Siberia. 

Myotis emarginatus Geoffroy, 1806 Geoffroy's Bat 

Approximate distribution of species: Holland, France, Germany, Switzerland, 
Italy, Hungary', Greece, Crimea, Transcaucasia, Russian Turkestan, Palestine, 
Persian Baluchistan. 

Myotis e.margin.^tus emargin.^tus E. Geoffroy, 1806 

1806. Vespertilio emarginatus Geoffroy, Ann. Mus. H.N. Paris, 8: 198. Charlemont, 

Givet, Ardennes, France. 
1844. Vespertilio rufescens Crespon, Faune Meridionale, /.• 20. Near Nimcs, Gard, 

France. Not of Brehm, 1829. 
1853. Vespertilio ciliatus Blasius, Arch. Naturgesch. 79, i : 287. Near Cologne, 

1856. Vespertilio sehrankii Kolenati, AUg. Dtsch. Naturh. Ztg. 2: 178, nom mid. Not of 

Wagner, 1843. 
1880. Myotis ciliata var. budapestiensis Margo, Magvar orv. es termeszetvisg. xx, 

nagygyiil. munk, 255. Budapest, Hungary. 
1890. Vespertilio neglectus Fatio, Arch. Sci. Geneve, 24: 512. Valavran, near Geneva, 


Range: Europe, as above, east to the Caucasus. 

Myotis em.a.rgin.'^tus desertorum Dobson, 1875 

1875. Vespertilio desertorum Dobson, in Blanford, Ann. Mag. N.H. 16: 309. Jalk, 

Persian Baluchistan. 
1920. Myotis lanceus Thomas, J. Bombay N.H. Soc. 26: 933, misprint, corrected to 

lanaceus Wroughton, 1920, J. Bombay N.H. Soc. 2y: 316. Dizak district, 

3,820 ft., Persian Baluchistan. 

Myotis em.\rginatus turcom.\nicus Bobrinskii, 1925 

1925. Myotis emarginatus turcomanicus Bobrinskii, Bull. Soc. Nat. Moscou, ^4: 358. 
Murgab Valley, Turkmen-Kala, Russian Turkestan. 

Myotis emarginatus saturatus Kuzyakin, 1934 

1934. Myotis lanaceus saturatus Kuzyakin, Bull. Soc. N.H. Moscou, ^j: 320, 329. 
Tashkent, Russian Turkestan. 



The two following named species seem allied to emarginatus. Tate placed the second 
in subgenus Mvotis, but the measurements he gives are too small for that subgenus. 

Myotis peytoni W'roughton & Ryley, 1913 

1913. Mxotis pntoni \\'roughton & Ryley, J. Bombay N.H. Soc. 22: 13. Gersoppa 
Falls, 1,300 ft., Kanara, Southern India. 

Myotis pri.mula Thomas, 1920 

1920. Mrolii primula Thomas, J. Bombay X.H. Soc. 2y: 248. Pashok, 3,500 ft., near 
Darjeeling, Xorth-Eastern India. 

Myotis altariutn Thomas, 191 1 

Approximate distribution of species: Szechuan, China; and has also (1949) been 
recorded from Kweichow, China. 

Myotis .\ltarium Thomas, 191 1 

1911. Mrolis altarium Thomas, Abstr. P.Z.S. 3; P.Z.S. 161. Omei Shan (Omi San), 
Szechuan, China. 

Myotis siligorensis Horsfield, 1855 

Approximate distribution of species; Kumaon, Nepal, Sikkim; Fukien, Southern 
China; Tonkin, Indo-China; Siam. 

Myotis siligorensis siligorensis Horsfield, 1855 

1855. Vespertilio siligorensis Horsfield, Ann. Mag. X.H. /6'.- 102. Siligori, Xepal. 

{W'roughton gave Darjeeling.) 
(?) 1855. Vespertilio darjilingensis Horsfield, loc. cit. 
Range: includes Kumaon, Sikkim. 

Myotis siligore.n'sis sowerbyi Howell, 1926 

1926. Mvotis sowerbyi Howell, Proc. Biol. Soc. Washington, 29- 'S^- Venpingfu, 
Fukien, 3,000 ft.. Southern China. G. Allen referred this to laniger as a 
synonym (which it is not, according to Tate and Osgood) and quoted that 
form from Yunnan, Fukien, Hainan. 

Myotis siligorensis alticraniatus Osgood, 1932 

1932. Myotis siligorensis alticraniatus Osgood, Field Mus. Publ. Zool 18: 232. Muong 
Moun, Tonkin, Indo-China. 

.Myotis siligorensis thaianus Shamel, 1942 

1942. Myotis siligorensis thaianus Shamel, J. Mamm. 2j: 323. Chiengmai, Siam. 

Myotis frater G. Allen, 1923 

Approximate distribution of species: Fukien, South-Eastern China; and most 
likely represented in Koiea, the Southern Ussuri district of Eastern Siberia, the 
Krasnoiarsk district fSiberia) and Tadjikistan I'Russian Turkestan). 



Myotis frater frater G. Allen, 1923 

1923. Alyotis frater G. Allen, Amer. Mus. Nov. No. 85: 6. Yenping, Fukien, South- 
Eastern China. 

Myotis (?) frater longicaudatus Ognev, 1927 

1927. Mvotis longicaudatus Ognev, J. Mamm. 8: 145. Vladivostock, Eastern Siberia. 

Range: to Korea, and the Siberian localities listed above. The published 

measurements are very similar to those o{ frater. 

Subgenus ISOTUS Kolenati, 1856 

Myotis nattereri Kuhl, 181 8 Natterer's Bat 

Approximate distribution of species: Britain, Ireland, France, Switzerland, Spain, 
Italy, Holland, Denmark, Sweden, Norway, Poland, Germany; Crimea, Caucasus, 
a few places in Russia (including near Leningrad, Kirov (formerly Vyatka) ) ; 
Kopetdag (South-\Vestern Turkestan), Sayan Mountains, Southern Yakutia, Amur 
Valley, east to Vladivostock; Japan, Korea and Manchuria (Kuzyakin). 

Myotis nattereri nattereri Kuhl, 18 18 

1818. Vespertilio nattereri Kuhl, Ann. Wetterau Ges. Naturk. ^, i : 33. Hanau, Hessen, 

1863. Isotus nattereri var. typus Koch, Jb. Nassau. Ver. Naturk. 18: 430. Wiesbaden, 

1863. Isotus nattereri var. spelaeus Koch, loc. cit. Erdbach, Nassau, Germany. 

1904. Myotis escalerai Cabrera, Mem. Soc. Esp. H.N. 2: 279. Foyos, near Valencia, 

Range: Europe. 

Myotis nattereri bombinus Thomas, 1905 

1905. Myotis nattereri bombinus Thomas, P.Z.S. igo§, 2: 337. Tano, Miyasaki Ken, 

Kiushiu, 500 ft., Japan. 

Myotis nattereri amurensis Ognev, 1927 

1927. Myotis nattereri amurensis Ognev, J. Mamm. 8: 144. Amur River, Eastern 
Siberia. Ranges to Northern Korea. 

Myotis nattereri tschuliensis Kuzyakin, 1935 

1935. Myotis nattereri tschuliensis Kuzyakin, Bull. Soc. Nat. Moscou, .^4: 434, 437. 
Tschuli (Chuli), Kopet-dag Mountains, South-\Vest Russian Turkestan. 

Subgenus PARAMYOTIS Bianchi, 1916 

Myoris bechsteini Kuhl, 1818 Bechstein's Bat 

Approximate distribution of species: England, France, Belgium, Holland, Spain, 
Switzerland, Germany, Hungary, Sweden, Poland, Lithuania, Ukraine, Caucasus 
and North-Western Transcaucasia 



Myotis bechsteini Kuhl, 181 8 

1818. Vespertilio bechstcinii Kuhl, Ann. Wcttcrau. Gcs. Naturk. ^, i: 30. Hanau, 

Hcsscn, Germany. 
1902. Vespertilio ghidinii Fatio, Rev. Suisse Zool. 10: 401. See also Fatio, 1905, Arch. 

Sci. Geneve, ig: 511. Lugano, Ticino, Switzerland. 
1906. Myotis bechstcinii favonicus Thomas, Ann. Mag. N.H. 18: 220. La Granja, 

northern side of Sierra de Guadarrama, Segovia, Spain. 

Subgenus MYOTIS Kaup, 1829 

We provisionally suggest that Myotis blythi (Tomes) is the first name for a species 
hitherto called oxvgnathiis which is much like M. myotis but occurs with it fairly 
extensively in Europe and averages smaller in size. From these two species M. 
siccirit/s scenes quite distinct, both cranially and dentally. 

Myotis myotis Burkhausen, 1797 Large Mouse-eared Bat 

Approximate distribution of species: France, Switzerland, Itah', Sardinia, Spain, 
Portugal, Germany, Poland, Hungary, Rumania; Southern Sweden, and has once 
been recorded from England. Eastwards to the Soviet Carpathians (Kuzyakin); 
Shensi, Szechuan, Yunnan, Chekiang and Fukien, China; Persia and Afghanistan. 

As here listed, this is equivalent to the largest members of Myotis [sensu stncto) as 
listed by Tate, 1941, p. 548. 

Myotis myotis myotis Borkhausen, i7f)7 

1774. ]'cspertilii> minimis Schreber, Saugeth. /; 1G5, and of Dobson, Blanfird, and 

earlier authors, but not of Linnaeus, 1758. 
1797. Vespertilio myotis Borkhausen, Deutsche Fauna, /.• 80. Thuringia, Germany. 
1797. Vespertilio tnyosotis (E.A.) Compend. Bibliothek, 21 ( Zoologe 5-1?) : 46. (This 

work does not appear to be available in London, and the reference is quoted 

as given by .Sherborn. Other authors have rjuoted it as of Borkhausen and as 

of Bechstein, with dates 1797 or 1800.) 
1827. Vesficrtdio siihmurimis Brehm, Ornis, j.- 23. Renthcndorf, Thuringia, Germany. 
1844. Vespertilio latipennu Crespon. Faune Meridionale, /.■ 17. Near .\imcs. Card, 

1863. Myotis nuiiiiiiis var. typiis Koch, Jb. Nassau \'er. Naturk. 18: 415. Wiesbaden, 

Nassau, Germany. 
1863. \tyotis murviu\ \ar. alpinus Koch, loc. cit. St. Gothard, Uri, Switzerland. 
1886. Myotis muniiii \pclaea Bielz, Verb. Mitt. SiebenbUrgischen Ver. Nalurw. 

Hermaiinst.idt, ^6'.- 83. Homorod-Almas Cave, Hungary .Nee Kiich, 1863. 

Range: Europe. 

Myoiis myotis {hinensis Tomes, 1857 

l8-,7. Ve\/iertilio ehiiien\i\ Fomcs, P.Z.S. 52. Soutliern C^hina. Range: \'unnau to 
1 ukien. 



Myotis myotis omari Thomas, 1906 

1906. Mrotis myotis omari Thomas, P.Z.S. igo^, 2: 521. Derbent, 50 miles west of 
Isfahan, 6,500 ft., Persia. (Ognev also recorded it from Kopet-Dag, South- 
West Russian Turkestan.) 

Myotis myotis ancilla Thomas, 1910 

1910. Myotis myosotis ancilla Thomas, Abstr. P.Z.S. 25; P.Z.S. 636. Shangchow, 
South-Eastern Shensi, China. 

Myotis myotis risorius Cheesman, 1921 

1 92 1. Myotis myotis risorius Cheesman, J. Bombay N.H. Soc. sy: 575. Shiraz, 5,200 ft. 

Myotis myotis luctuosus G. Allen, 1923 

1923. Myotis chinensis luctuosus G. Allen, Amer. Mus. Nov. No. 85: 5. Wanhsien, 
Szechuan, China. 

Myotis blythi Tomes, 1857 

Appro.ximate distribution of species, as here understood: Spain, Switzerland, 
Austria, Italy, Sardinia, Malta, Montenegro, Greece, Crete; U.S.S.R. localities in- 
clude Moldavia, Crimea, Caucasus, Turkmenia, ^Veste^n Tianshan, Hissar-Alai 
Mountains and Turanskaya Lowlands. Kuldja (Western Chinese Turkestan) 
according to Ognev. Rajputana, Punjab and perhaps Kashmir. Asia Minor and 
Palestine (according to Kuzyakin, in Bobrinskii). Algeria, Tunis, Morocco. 

Myotis blythi blythi Tomes, 1857 

1857. Vespertilio blythii Tomes, P.Z.S. 53. Nasirabad, Rajputana, India. Ranges to 
Simla, Northern India. 

Myotis (?) blythi dobsoni Trouessart, 1878 

1873. Vespertilio murinoides Dobson, J. Asiat. Soc. Bengal, ^2, 2: 205. Not of Lartct, 

1 85 1. Chamba, 3,000 ft., North- Western Himalayas. 
1878. Vespertilio dobsoni Trouessart, Rev. Zool. Paris, 6: 248. New name for murinoides 

Dobson, preoccupied. Synonym oi blythii, according to Wroughton. 

Blanford listed the form Vespertilio africanus Dobson, 1875, '^i synonymy with blythi, 
but it is thought to have come from Gabon, \Vest Africa. (See G. M. Allen, 1939, 
Checklist African Mammals.) 

Myotis (?) blythi oxygnathus Monticelli, 1885 

1885. Vespertilio oxygnathus Monticelli, Ann. Accad. Aspir. Nat. /; 82. Matcra, 
Basilicata, Italy. 

Range: Europe, Turkestan, North-\Vest Africa and South-\Vestern Asia, as listed 



Myotis sicarius Thomas, 191 5 

Approximate distribution of species: Sikkim, India. 

Myotis sicarius Thomas, 1915 

1915. Mwtis sicarius Thomas, J. Bombay N.H. Soc. 23: 608. Northern Sikkim. 

Subgenus CHRTSOPTERON ]cnUnk, 1910 

Myotis formosus Hodgson, 1835 Hodgson's Bat 

Approximate distribution of species: Fukien and adjacent states in Southern 
China, Nepal, Punjab, Kumaon (Blanford also quoted it from Sikkim, Bengal, 
Assam); Korea, Formosa, Southern Japan. 

Myotis formosus formosus Hodgson, 1835 

1835. Vespertilio formosa Hodgson, J. Asiat. Soc. Bengal, 4: 700. Nepal. Range in- 
cludes Kumaon and Punjab. 

1863. Kcrivoula pallida Blyth, Cat. Mamm. Mus. Asiat. Soc. Bengal, 34. Chaibassa, 
Orissa, India. 

1 87 1. Vesper lilio auratus Dobson, J. Asiat. Soc. Bengal, 40, 2: 186. Darjeeling, India. 

Myotis formosus rufoniger Tomes, 1858 

1858. Vespertilio rufo-niger Tomes, P.Z.S. 79, pi. 60. Shanghai, Kiangsu, China. 
Range includes Fukien, China. 

Myotis (?) formosus andersoni Trouessart, 1897 

1881. Vespertilio dobsoni Anderson, Cat. Mamm. Ind. Mus. 143. Purneah, Bengal. 
Not of Trouessart, 1878. 

1897. Vespertilio andersoni Trouessart, Cat. Mamm. 129. New name for dobsoni Ander- 
son, preoccupied. 

Myotis formosus tsuensis Kuroda, 1922 

1922. Myotis tsuensis Kuroda, J. Mamm. 3: 43. Izugahara, Tsushima Island, Southern 
Japan. iStatus/(/c Kuroda.) 

Myotis formosus watasei Kishida, 1924 

1924. Myotis watasei Kishida, Zool. Mag. Tokyo, 36: 30-49, 127-139. {M.V.). 
Terason, Formosa. 

Myotis formosus chofusukei Mori, 1928 

1928. Mvolis chofusukei Mori, Annot. Zool. Jap. //.■ 389. Kaishu, Kokaido, Korea. 

Subgenus LEUCONOE Boie, 1830 

Tate (194 1, 550) divides this subgenus into five sections, typified by daubentoni, 
capaccinii, davidu adversus and dasycneme. There are several other standing species. Of 



these, A/, peqiiinius is from descriptions certainly valid. Another early name, M. macro- 
dactylus, is regarded as a subspecies of M. capaccinii by Kuzyakin, in Bobrinskii (1944), 
but as noted by Thomas (1906, P-Z-S. 1903, 2: 337) this is an error. Mr. R. W. 
Hayman states that in the extensive series in the British Museum the tibia and 
adjacent membrane are not furred, thereby differing from capaccinii. Tate placed the 
species tentatively in his adversus section, and Mr. Hayman states M. macrodactylus 
differs from M. adversus and M. daubentoni by the attachment of the wing membrane, 
which is high on the tibia in macrodactylus, not so in the other two species just 

Myotis daubentoni Kuhl, 18 19 Daubenton's Bat. Water Bat 

Approximate distribution of species: Britain, Ireland, Norway, Sweden, France, 
Switzerland, Holland, Denmark, Spain, Italy, Germany, Rumania, Poland; Russia 
and Siberia, eastwards to Kamtchatka, Sakhalin and Ussuri region, its northern limit 
runs close to the Goth parallel, and its southern limit from Southern Ukraine, 
Southern Volga, Northern Kazakstan, the Altai. Tate quotes it from Japan and the 
Kurile Islands. Manchuria, Mongolia, Fukien (? Yunnan and Hainan), China; 
Bodenheimer quotes it from Palestine. 

Myotis daubentoni daubentoni Kuhl, 18 19 

1819. Vespertilio daubentonii Kuh. Ann. Wetterau. Gcs. Naturk. ./, 2: 195. Hanau, 
Hessen-Nassau, Germany. 

1839. Vespertilio aedilis jenyns, Ann. Nat. Hist, j: 73. Aukland St. Andrew, Durham, 

1844. Vespertilio lanatus Crespon, Faune Meridional. /.• 15. South of Nimes, Gard, 

1 87 1. Vespertilio capucinellus Fitzinger, S.B. Akad. Wiss. Wien, 6j, i : 206. ? Bavaria. 
1 87 1. Vespertilio minutellus Fitzinger, loc. cit. ? Bavaria. 

1871. Vespertilio daubentonii albus Fitzinger, loc. cit. 210. Renaming oi aedilis ^enym. 
1890. Vespertilio staufferi Fatio, Faune Vert. Suisse, j, 3me suppl. aux Mamm. 6. 

Lucerne, Switzerland. 
Range: Europe. 

Myotis daubentoni volgensis Eversmann, 1840 

1840. Vespertilio volgensis Eversmann, Bull. Soc. Nat. Moscou, 24. Ural Mountains, 

Eastern Russia. 
1912. Myotis petax Hollister, Smiths Misc. Coll. 60: 6. Kosh-Agatch, Chuiskaya 
steppe, 7,300 ft., Altai district, Siberia. 

Myotis (?) daubentoni laniger Peters, 1871 

1 87 1. Vespertilio laniger Peters, in Swinhoe, P.Z.S. i8yo: 617. Amoy, Fukien, China. 

Myotis daubentoni ussuriensis Ognev, 1927 

1927. Myotis daubentonii ussuriensis Ognev, J. Mamm. 8: 146. Near Vladivostock, 
Eastern Siberia. Ranges to Sakhalin, Korea. 



Myotis daubenton'i loukashkini Shamel, 1942 

10)42. Myotis pelax loukashkini Sham(?l, Proc. Biol. Soc. \\'ashiiit;ton, jj.' 103. 
Wutaliciu liich. Third Lake, Hcilunt;kiano- Province, Xorllicrn Manchuria. 

Myotis capaccinii lionapartc, 1837 Long-fine;crcd Bat 

Approximate distribution of species: Southern France, Spain, Italy, Switzerland, 
.Sardinia, Transylvania, Bulgaria; Lower Amu-Darya, Russian Turkestan; besides 
this Kuzyakin cjuotcs the species from the southern Maritime Province of Siberia 
(under the name .\/. c. macrodaclvlus; but macrodactvliis is not capaccinii, see above under 
subgenus Lcnconoc); Morocco, Algeria. 

.Myotis cap.\cci.\ii capaccinii Bonaparte, 1837 

1837. Vespeiiilio capaccinii Bonaparte, Faun. Ital. /, fasc. 20. Sicily. 

1840. Vespertilto mcgapodms Temminck, Mon. Mamm. .'.• 189. Sardinia. 

1 84 1. Vcspertilio dasypus de Selys Longchamps, Atti dclla scconda Riun. dcgli Sci. 

Italiani, Torino, 1840: 247. Sardinia. 
(?) 1844. Vespeiiilio pelluccns Crespon, Faune Meridionalc, /.• 16. Cave near Pont- 

du-Gard, Gard, France. 
(.') i860. Brachyotus hlaui Kolcnati, Jh. Miihr. Schl. Ges. Ackerbau, i8^c): 102. 

Swabia, Southern Bavaria, Germany. 
1878. Vcspertilio majori Xinni, Atti R. 1st. Veneto, 4, i: 721. Substitute for blasii 

Forsyth .Nlajor, 1877, Atti Soc. Tosc. Sci. Nat. Pisa, jj.- 108. 
Range: Europe, Morocc(j and Algeria. 

Myotis cAP;\cciNn burf.schi Heinrich, 1936 

1936. Leuconoe capaccinii bureschi Heinrich, Mitt. Naturw. Inst. Sofia, 9.' 38. Karamler, 
StraiidjaT5alkan, 800 ft., Bulgaria. 

The two following-named species are allied to M. capaccinii, and possibly represent it. 

Myotis fimbriaius Peters, 1871 

1 87 1. Vespertilio Jimhriatus Peters, P.Z.S. l8jn: G17. Amoy, Fukien, China. 

1926. Myntis hirsutns Howell, Proc. Biol. Soc. Washington, ^9.- 139. ^'enpingfu, 
Fukien, 2,000 ft., China. 

.Myotis lo.n'cipfs Dobson, 1873 

(?) 1855. -Uj'f'''* theobaldi Blyth, J. Asiat. Soc. Bengal, r^.- 363. Caves near Matar 

Xag, north of Islamabad, Kashmir. Thomas, 1915, J. Bombay N.H. Soc. 

-",•; (no, agues with Blanford lliis form should be considered un- 


1872. I'espeititio macropus Dobson, Proc. Asiat. Soc. Bengal, 209. Not of Gould, 1854. 

Caves of Bhima Devi, 6,000 ft., Kashmir. 

1873. Ve^pertilm lon<;ipes Dobson, Proc. .'Xsiat. Soc. Bengal, 1 10. Renaming nl' macropus, 

'?,! 187-,. I'npntilio meoalopus Dobson, Ann. -Mag. .N.H. 16: 261. ? Kashmir. Dobson 
ga\e the loiality as Gaboon, West Africa, but Thomas, 1915,,!. Bombay 
N.H. Soc. ^"j.- 610, said the type was identical with a cotype (A'loni^ipes and 
certainly did not come from Gaboon. 



Myotis pequinius Thomas, 1908 

Approximate distribution of species: Chihii, China. 

Myotis pequinius Thomas, 1908 

1908. Afyotis (Leuconoe) pequinius Thomas, P.Z.S. 637. Thirty miles west of Pekin, 
ChihH, 600 ft., China. 

Myotis davidi Peters, 1869 

Approximate distribution of species: Chihii, and apparently Hainan and Kiangsi, 

Myotis davidi Peters, 1869 

1869. Vespertilio davidii Peters, Mber. Preuss. Akad. Wiss. 402. Pekin, Chihii, China. 

Myotis adversus Horsfield, 1824 

Approximate distribution of species: Malay States, Java, probably Borneo, 
Sumatra, ? Celebes, ? Australia; for status of type specimens and immediate allies, 
see Tate, 1941, Bull. Amer. Mus. N.H. j8: 551. Siam. ? Ceylon (the form quoted from 
Ceylon by Wroughton (1918) as "hasselti", forearm 40 mm. in the key, cannot be 
hasselti, as Tate shows this to have been based on a small form, with forearm 32 mm.). 
Possibly also represented in Formosa, Tibet, the Andaman Islands and Southern 

The listing of this species is provisional. 

(Myotis adversus adversus Horsfield, 1824. Extralimital) 
1824. Vespertilio adversus Horsfield, Zool. Res. Java, (8). Java. 

Myotis (?) adversus dryas Andersen, 1907 

1907. Myotis dryas Andersen, Ann. Mus. Stor. Nat. Genova, 5.- 33. Port Blair, South 

Andaman Islands, Bay of Bengal. 

Myotis (?) adversus taiwanensis Arnbiick-Christie-Linde, 1908 

1908. Aiyotis taiwanensis Arnbiick-Christie-Linde, Ann. Mag. N.H. 2: 235. Takao, 

Anping, Tainan, Formosa. Range: has also been recorded from Tibet. 

Myotis (?) adversus peshwa Thomas, 1915 

1915. Leuconoe peshwa Thomas, J. Bombay N.H. Soc. 2^: 61 1. Poona, Bombay, India. 

Myotis adversus continentis Shamel, 1942 

1942. Myotis adversus continentis Shamel, J. Mamm. 23: 323. Bangkok, Siam. 

Myotis (?) adversus (?) subsp. 

1918. Leuconoe hasselti Wroughton, J. Bombay N.H. Soc. 25.- 598. (Not hasselti Tem- 

minck, 1840, from Java.) Ceylon, quoted by Wroughton from Northern, 

Central and Eastern Provinces. 


PALAF,ARC:TIC: and IMJIAX mammals 1758-194G 

Tate thought the ftillowing form should be referred to the adversus section of 
Leuconoe, but Chaworth-Mustcrs, in a paper he was preparing on the bats of Arabia, 
shortly before his death, made the suggestion that dogalensis was based on a young 
specimen of the African Myotis bocagei Peters, 1870, which Tate (p. 552) refers to 
subgenus Selysius. 

Myotis dogalensis Monticelli, 1887 

1887. Vespertilio dogalensis Monticelli, Ann. Mus. Stor. Nat. Genova, j.- 518. Aden, 

Myotis macrodactylus Temminck, 1840 
Approximate distribution of species: Japan. 

Myotis m.^crodactylus Temminck, 1840 

1840. Vespertilio macrodactylus Temminck, Mon. Mamm. 2: 231, pi. 58, figs. 3, 4, 5. 
Japan. Known from South Kuriles, Hondo, Shikoku, Kiushiu. 

Myotis dasycneme Boic, 1825 Pond Bat 

Appro.ximate distribution of species: Holland, Belgium, Northern France (Kuzya- 
kin), Denmark, Sweden, Poland; Russia, between 49" and 60" N., eastwards across 
^Vestern Siberia to the Yenesei. 

Myotis dasyc.n' dasycneme Boie, 1825 

1823. Vespertilio mystacinus Boie, Isis, Jena, 965, not of Kuhl, 1819. 

1825. Vespertilio dasycneme Boie, Isis, Jena, 1200. Dagbieg, near Wiborg, Jutland, 

Denmark. Renaming oi mystacinus Boie, preoccupied. 
1839. Vespertilio limnophilus Temminck, Mon. Mamm. 2: 17G, pi. 48, figs, i, 2. 


Myotis dasycneme m.'^jor Ognev & Worobiev, 1923 

1923. Myotis dasycneme major Ognev & Worobiev, Fauna Terr. Vert. Govt. Voronesh, 
98. Voronesh, Russia. 

Subgenus RICKETTIA Bianchi, 1916 

Myotis ricketti Thomas, 1894 Rickctt's Big-footed Bat 

.Approximate distribution of species: China, states of Fukien, .\nhwei. Shantung. 

Myotis rk;ketti Thomas, 1894 

(?) 1869. Vespertilio {Leuconoe) pilosa Peters, Mber. Preuss. Akad. \Viss. 403. Thought 

to be from Uruguay, South America. G. Allen (1938, 224) uses this name, 

but is not followed by Tate (1941). 
1894. Vespertilio {Leuconoe) ricketti Thomas, Ann. Mag. N.H. 14: 300. Foochow, 

Fukien, China. 


Incertae sedis 

1863. Myotis ? berdmorei Blyth, Cat. Mamm. Mus. Asiat. Soc. 35, based on descrip- 
tion without name in Blyth, 1859, J. Asiat. Soc. Bengal, 28: 293, of a bat 
from Schwegyin, Burma. ? Unidentifiable; see Blanford, 1891, Fauna Brit. 
India, Mamm. 330. 

1942. Myotis deignani Shamel, J. Mamm. 2j.- 324. Chiengmai, Siam, Tate (1947, 

Mamm. E. Asia) lists it in subgenus Selysius. 

1943. Myotis coluotus Kostron, Rozpr. Ceske Akad. 52, 17: i, and Bull. Int. Acad. 

Prague, ^2- '90- Jaworzitschko, Northern Moravia, Czechoslovakia. 

1944. Myotis flavus Shamel, J. Mamm. 25.- 191. Enri, Formosa. Tate (1947, Mamm. 

E. Asia) lists it in subgenus Chrysopteron. 
1944. Myotis abei Yoshikura, Zool. Mag. Tokyo, ^6 (i, 2, 3): 6. {N.V.). Southern 
Sakhalin. (In Japanese, but title is "On a new Whiskered Bat".) 

1 93 1. Pactia mori Kishida & Mori, Zool. Mag. Tokyo, 43: 378, Korea, nom. nud. 

Genus DISCOPUS Osgood, 1932 

1932. Discopus Osgood, Field Mus. Publ. Zool. 18: 236. Discopus denticulus Osgood. 

1 species : Discopus denticulus, page 1 5 1 

Discopus denticulus Osgood, 1932 

Appro.ximate distribution of species: Laos, in Indo-China. 

Discopus denticulus Osgood, 1932 

1932. Discopus denticulus Osgood, Field Mus. Publ. Zool. 18: 236. Phong Saly, Laos, 

Genus VESPERTILIO Linnaeus, 1758 

1758. Vespertilio Linnaeus, Syst. Nat. loth ed. /.■ 31. Vespertilio murinus Linnaeus. 
1839. Vesperugo Keyserling & Blasius, Arch. Naturg. 5, i : 312. Contained 13 species, 

one of which was discolor = murinus. 
1839. Vesperus Keyserling & Blasius, loc. cit. 313 (part). Not of Latreille, 1829. 
1856. Meteorus Kolenati, Allg. Dtsch. Naturh. Ztg. 2: 131 (part). (Included seyeral 

species, one of which was discolor = murinus.) 
1863. Aristippe Kolenati, Horae Soc. Ent. Ross. 2, 2: 40 (part; included murinus). 
1872. Marsipolaemus Peters, Mber. Preuss. Akad. Wiss. 260. Vesperugo albigularis 

Peters = Vespertilio murinus Linnaeus. 

2 species: Vespertilio murinus, page 152 

Vespertilio superans, page 152 

On this genus and all genera of Vespertilioninae except Myotis, see Tate, 1942, 
Bull. Amer. Mus. N.H. 80: 221-297. As restricted by Miller, this genus contains two 
species, murinus and superans. Kuzyakin, in Bobrinskii, 1940, Mamm. U.S.S.R., shows 
clearly that both these forms are valid species. This author refers Eptesicus and 
Pipistrellus to the present genus (but keeps Nyctalus separate). The cranial and ear 


i'ai..\i;arc:tic and Indian mammals ]7-,8-i946 

details noted by Miller as restricting the genus to the present species arc not perhaps 
of great importance, but the two allied genera referred to above arc both so widely 
distributed and contain so many species that it is a matter of coineniencc to retain 

Vespertilio murinus Linnaeus, 1 758 Particoloured Bat 

Approximate distribution of species: Norway, Sweden, Denmark, France, Ger- 
many, Clzechoslovakia, Poland, Switzerland, Austria. Has been very rarely taken in 
England ("a single specimen, utidoubtcdly a straggler, taken at Plymouth" (Miller, 
1912) and has more recently been recorded from the Shetland Islands (Ritchie, 1927, 
Scot. Nat. Edinburgh, 'loi) ). Russia, from about 60 N., south to the Black Sea and 
Caucasus, Russian Turkestan, and across Siberia to the Ussuri district. Japan; 
Mongolia; Kashmir; Persia; Kashgar (Chinese Turkestan). 

Bodenhcimer quoted \ . murinus from Palestine, but this is far from the normal 
range of the species, and it must be borne in mind that in earlier literature .Mrolis 
mrotis, which occurs in South-Western Asia, used to be called "IVi/x/V/Z/o murinus". 

Vespertilio muri.\us muri.nus Linnaeus, 1758 

1758. Vcsjifrtilio murinus Linnaeus, Syst. Nat. loth cd. /.■ ;v:!. Sweden. 

l8ir). Vespertilio (lisenlnr N.iltcrcr in Kulil, Ann. Wctterau. (ics. Naturk. 4, 2: 187. 

\'ienna, Austria. 
i8y5. Vesperugo krascheninnikovi Evcrsmann, Bull. Soc. Nat. Moscou, 36, 2: 488. 

Orenburg, Russia. 
1872. I'esperus [Marsipolaemus) alhigukiris Peters, Mbcr. Preuss. Akad. Wiss. 260. 

(Type supposed to have been taken in Mexico. See Miller, 1912, Cat. 

Mamm. \V. F.uropc, 238.) 
1885. Vesperus siculus Daday, Orv. Term. Ert. Koloszwar, 10: 275. Homorod-Almas 

Clave, Hungary. 
lOov Vespertilio discolor luleiis Kastschcnko, Trans. Tomsk. Lhiiw .7.- i02d. Ner- 
chinsk, Transbaikalia, Eastern Siberia. 
if)i3. ]'e\/iertilio discolor michnoi Kastschenko, Annu. Mus. Zool. Acad. St. Petersb. 

ty: 391. Aga, Aginska Steppe, Transbaikalia, Eastern Siberia. 
Range: as in the species, except Japan. 

Vespertilio ,?)mi'ri\ls x.wiivei Kuroda, 1920 

1920. Mvctalus nocluln namiyei Kuroda, Annot. Zool. Jap. (), 5: 601. Otsukuejima, 
coast of C'hikuzen Province, Kiushiu, Japan. 

Vespertilio superans Thomas, 1899 

Approximate distribution of species: Maritime Province of Eastern Siberia; 
Hokkaido and Hondo: Korea; China, states of Szcchu.m, Fukien, Shansi, Chihli, 
? Kansu; and .MoMi^olia. 

Vesper iiLiu siiplra.ns Thomas, 1899 

1899. Vespertilio murinus superans Thomds, P.Z.S. i8()8: 770. Sesalin, Ichang, Hupeh, 

I =,2 


Dobson (1878) (followed by Blanford, 1891, Mamm. Brit. India) called the genus 
now known as Mvotis by the name Vespertilio, and the present genus, in a much wider 
sense than as accepted by Miller, "Vesperugo" . V. murinus was called ''Vesperugo dis- 
color", and V. murinus of Dobson is the species now known as Alyotis myolis. 

Genus EPTESICUS Rafinesque, 1820 

1B20. Eptesicus Rafinesque, Annals of Nature, 2. Eptesicus mdanops Rafinesque = 

Vespertilio fuscus Beauvois, from North America. 
1829. Cnephaeus Kaup, Skizz. Europ. Thierw. i: 103. Vespertilio serotinus Schreber. 
1837. Noctula Bonaparte, Faun. Ital. /.■ fasc. xxi. Noctula serotina. 
1856. Cateorus Kolenati, AUg. Dtsch. Naturh. Ztg. 2: 131. Vespertilio serotinus Schrcher. 
1858. Amblyotus Kolenati, S.B. Akad. Wiss. Wien, 2g: 252. Amhlyotus atratus Kolenati 

^ Vespertilio nilssonii Keyserling & Blasius. 
1866. Pachvomus Gray, Ann. Mag. N.H. jy: 90. Scotophilus pachyomus Tomes. 
1870. Nrctiptenus Fitzingcr, S.B. Akad. Wiss. Wien, 62: 424. Vespertilio smithii Wagner, 

from South Africa. 
1892. Adelonycteris H. .Allen, Proc. Acad. Nat. Sci. Philadelphia, 1891 : 466 (part). 

(Substitute for Vesperus Keyserling & Blasius, 1839, which is preoccupied 

by Vesperus Latreille, 1829, and contained species of both the present genus 

and Vespertilio.) 
191 7. Pareptesicus Bianchi, .-\nnu. Mus. Zool. Acad. St. Petersb. 21: lxx\i. Vesperugo 

pachyotis Dobson. 
1917. Rhyneptesicus Bianchi, Annu. Mus. Zool. Acad. St. Petersb. 21: lxx\i. Vesperugo 

nasutus Dobson. 
1926. Neoromicia Roberts, Ann. Transvaal Mus. //.• 245. Eptesieus z'tluensis Roberts, 

from Natal. 
1931. Tuitatus Kishida & Mori, Zool. Mag. Tokyo, 43: 372-391 [N.V.) nom. nud. 
1934. Vespadelus Iredale & Troughton, Mem. Austr. Mus. 6: 95. Australian species 

oi Eptesicus. (Nom. nud.) 

This genus is nearly world-wide. It is near Vespertilio, and referred to that genus by 
Kuzyakin. It is not easy to say how many species there are in the present region, but 
the following seven seem certainly valid : 

Eptesicus isabellinus, page 156 
Eptesicus nasutus, page 154 
Eptesicus nilssoni, page 1 55 
Eptesicus pachyotis, page 155 
Eptesicus serotinus, page 156 
Eptesicus sodalis, page 1 56 
Eptesicus walli, page 154 

Several subgeneric names are available, but we are inclined to ignore them until 
more detailed revision has taken place in the genus. Chaworth- Musters, in a key to 
Arabian bats which he was preparing shortly before his death, suggests that matschiei 
represents the Indian nasutus; this is accepted. The Turkestan form bobrinskoi is 
apparently approximately the same size, and has yet to be proved specifically distinct 


PALAEARtrnC: and INDIAN MAMMALS 1758-1946 

from nasutus. Kuzyakin regards the form ognevi as a valid species, but according to 
Ognev and Tate it is a race oi sodalis. We have provisionally united the forms innesi, 
isabdlinus and hottae (forearm about 40-44 mm.) under the prior name isahellimis. 
G. Allen listed isabellinus as a race oi serotinus, but according to Tate's measurements 
( 1942, 275), it is too small for that species. According to Kuzyakin (1944) some species 
formerly referred to this species should be transferred to Pipislrellus savii. 
For review, see Tate (1942, 271). 

EptesH'us nasutus group 
Rhxneptesicus Bianch! is available if subgeneric division is required. 

Eptesicus nasutus Dubson, 1877 Sind Bat 

Approximate distribution of species: Sind and Puirjab; Arabia, Persia; \i hohrinskoi 
is the same, deserts of Kazakstan and Russian Turkestan, Xorthcrn Osetia (? Cau- 
casus) and Yakutsk, Siberia. 

Eptesicus nasutus nasutus Dobson, 1877 

1877. Vesperugo [Vesperus) nasutus Dobson, J. Asiat. Soc. Bengal, .^6', 2 : 31 1. Shikarpur, 
Sind, Western India. Range includes Punjab. 

Eptesicus nasutus matschiei Thomas, 1905 

1905. Vespertilio matschiei Thomas, Ann. Mag. N.H. 16: 573. Jimcl, near Aden, 
850 m.. Southern Arabia. 

Eptesicus nasutus pellucens Thomas, 1906 

igo6. Vespertilio matschiei pellucens Thomas, P.Z.S. /505, 2 : 520. Ahwaz, Karun River, 
220 ft., South-VVestern Persia. 

Eptesicus (?) nasutus bobrinskoi Kuzyakin, 1935 

1935. Eptesicus bobrinskoi Kuzyakin, Bull. Soc. Nat. Moscou, 44: 435-437- Tjulek 

wells in Aral Kara-Kum (desert), 65 km. east of city of Aralskoje More, 

Russian Turkestan. 

Eptesicus walli group 
For note on cranial characters of this species, see Tate (1942, 274). 

Eptesicus walli Thomas, 19 19 Wall's Serotine 

.Approximate distribution of species: Iraq. 

Eptesicus \v,\lli Thomas, 191 9 

1919. Eptesicus walli Thomas, J. Bombay N.H. Soc. 26: 746. Basra, Iraq. 

Eptesicus puchyotts grfjup 
Pareptesicus Bianchi is available here if subgeneric division is required. 



Eptesicus pachyotis Dobson, 1871 Thick-eared Bat 

Approximate distribution of species: Assam. 

Eptesicus pachyotis Dobson, 1871 

1 87 1 . Vesperugo ( Vesperus) pachyotis Dobson, Proc. Asiat. Soc. Bengal, 211. Khasi Hills, 

Eptesicus nilssoni group 
The name Arnblyotus Kolenati is available if subgeneric division is required. 

Eptesicus nilssoni Keyserling & Blasius, 1839 Northern Bat 

Approximate distribution of species: Norway, Sweden, Germany, France, Switzer- 
land, Denmark, Czechoslovakia, Northern Italy, Poland; Russia, north to Kola 
Peninsula; in Siberia, the northern limit drops roughly to the 6oth parallel, and 
ranges east to the Pacific; southwards, in summer, to north of Moscow and Gorki 
Provinces, but in the autumn has been found in districts further south fNorthern 
Volga, Smolensk, Eastern Carpathians) (Kuzyakin, in Bobrinskii). It occurs in 
Russian Turkestan. Mongolia, perhaps Manchuria, Chinese Turkestan, Korea, 
Tibet; Kashmir. 

Eptesicus nilssoni nilssoni Keyserling & Blasius, 1839 

1836. Vespertilio kuhlii Nilsson, Ilium. Fig. Skand. Fauna, pt. 17, pi. 34, upper fig. 
Not of Kuhl, 1819. 

1838. Vespertilio borealis Nilsson, Ilium. Fig. Skand. Fauna, pt. 19, pi. 36, upper fig. 

Not of MuUer, 1776. 

1839. Vespertilio nilssonii Keyserling & Blasius, Arch. Naturgesch. 5, 1:315. Sweden. 
1858. Arnblyotus atratus Kolenati, S.B. Akad. Wiss. Wien, sg: 252. Altvater, 2,400- 

4,600 ft., Austrian Silesia. 
Range: Europe, Siberia to the Pacific, Gilgit (Kashmir). 

Eptesicus nilssoni gobiensis Bobrinskii, 1926 

1926. Eptesicus nilssonii gobiensis Bobrinskii, C.R. Acad. Sci. U.R.S.S., A, 96. 

Burchasteitala, Gobi Altai Mountains, Mongolia. Ranges into Russian 

Central Asia. 

Eptesicus nilssoni centrasiaticus Bobrinskii, 1926 

1926. Eptesicus nilssonii centrasiaticus Bobrinskii, C.R. Acad. Sci. U.R.S.S., A, 96. 
Ushchele Khatu, near Russk, Orin-Nor, Tibet. 

Eptesicus nilssoni kashg.aricus Bobrinskii, 1926 

1926. Eptesicus nilssonii kashgaricus Bobrinskii, C.R. Acad. Sci. U.R.S.S., A, 97. 
Khotan-Tagh, mountains of Russki, near Kashgar, Chinese Turkestan. 

Eptesicus nilssoni parvus Kishida, 1932 

1932. Eptesicus parvus Kishida, Lansania, Tokyo, 4, 31: 2. North Korea. {NA'.) 

Tate (1942) lists several forms (not seen by him) as races oi nilssoni which are here, 
following Kuzyakin in Bobrinskii, transferred to Pipistrellus savii. 

L 155 


Eplesiais serotinus group 

(The type species belongs here.) 

Eptesicus sodalls Barrctt-Hamihon, 1910 

Approximate distribution of species: Rumania, Switzerhmd, Russian Turkestan, 
where widely distributed in the south and east, to South-Western Mongolia (Kuzya- 
kin"! ; Iraq. 

Eptesicus sodalis sod.\lis Barrett-Hamilton, 1910 

1910. Vespcrtilio sodatis Barrett-Hamilton, Ann. Mag. N.H. f}: 291. Bustenari, Pra- 
hova, 840 m., in Carpathians, Rumania. 

Eptesicus sodalis ognevi Bobrinskii, 1918 

1918. Eptesicus ognevi Bobrinskii, Fauna & Flora of Russia, 13: 12. {M.V.) Bokhara 

district, Russian Turkestan. 

Eptesicus sodalis hingstoni Thomas, 1919 

1919. Eptesicus hingstoni Thomas, J. Bombay N.H. Soc. 26: 745. Baghdad, Iraq. 

Eptesicus isabellinus Tcmminck, 1840 

Approximate distribution of species: Libya, Egypt, Arabia. 

Eptesicus isabellinus isabellinus Temminck, 1840 

1840. VesperlUio isaltelliniis Temminck, Mon. Mamm. 2: 205, pi. 52, figs, i, 2. 

Environs of Tripoli, Libya. G. Allen listed this as a race of serotinus, but 

Tate's measurements make it too small for that. 

Eptesicus isabellinus bottae Peters, 1869 

1869. Vesperus Iwtlae Peters, Mber. Prcuss. Akad. ^Viss. 406. Yemen, Arabia. 

Eptesicus isabellinus innesi Lataste, 1887 

1887. Vesperugo (Vesperus) innesi Lataste, Ann. Mus. Stor. Nat. Genova, 4: 625, 2 te.xt 
figs. Cairo, Egypt. 

Eptesicus serotinus .Schreber, 1774 Seroline 

Apprdximate distribution of species: England, France, .Switzerland, Spain, Italy, 
Sardinia, Germany, Holland, Denmark, Hungary, Yugoslavia, Rumania, Greece, 
Poland; Russia and Siberia, where the northern limit runs through Kharkov and 
Orenburg, roughly eastwards to Lake Balkash, and southwards to the Caucasus and 
Russian Turkestan; Persia, Asia Minor, Palestine (Bodenheimer) ; Chinese Turkestan, 
Mongolia. Korea; Shensi, Shantung and Chihli, in China, also Yunnan, Fukien and 
Chekiang if fl/n/cMo«i is regarded as a representative; Kashmir, Rajputana; West 
Africa (part). 

I V 


Eptesicus serotinus serotinus Schreber, 1774 

1774. Vespertilio serotinus Schreber, Saugeth. /; pi. 53 (text, p. 167). France. 

1776. Vespertilio serotine Muller, Natursyst. Suppl. Regist. Band, 16. 

1827. Vespertilio wiedii Brehm, Ornis, j: 24. Renthendorf, Thuringia, Germany. 

1827. Vespertilio okenii Brehm, loc. cit. 25. Renthendorf, Thuringia, Germany. 

1844. Vespertilio incisivus Crespon, Faune Meridionale, /.• 26. Nimes, Gard, France. 

1B63. Cateorus serotinus typus Koch, Jb. Nassau. Ver. Naturk. 18: 466. Wiesbaden, 

Nassau, Germany. 
1863. Cateorus serotinus var. mfescens Koch, loc. cit. Freiburg, Breisgau, Germany. 
1885. Vespertilio serotinus var. transylvanus Daday, Orv. Term. £rt. Koloszvar, 10: 275. 

Also-Szocs, Szolnok-Doboka, Hungary. 
1904. Vespertilio serotinus insularis Cabrera, Mem. Soc. Esp. H.N. 2: 263. Minorca, 

Balearic Islands. 
1904. Vespertilio isahellinus Cabrera, Mem. Soc. Esp. H.N. 2: 264. Andalusia, 

Southern Spain. Not of Temminck, 1840. 
1904. Vespertilio boscai Cabrera, Mem. Soc. Esp. H.N. 2: 265. Muchamiel, Alicante, 

Range: Europe. 

Eptesicus serotinus turcomanus Eversmann, 1840 

1840. Vespertilio turcomanus Eversmann, Bull. Soc. Nat. Moscou, 21. Between Caspian 

and Aral Seas, Russian Turkestan. 
(?) 1865. Vespertilio (Vesperus) mirza de Filippi, Viagg. in Persia, 342. Persia. 
1875. Vesperugo albescens Karelin, Trans. St. Petersb. Nat. Soc. 6: 265, nom. nud. 

Sluda, near Gureva (? = Guriev, mouth of River Ural). 
Range: Russian Asia and Persia. 

Eptesicus serotinus pachyomus Tomes, 1857 

1857. Scotophilus pachyomus Tomti, P.Z.S. 50. Rajputana, India. Ranges to Kashmir. 

Eptesicus serotinus shiraziensis Dobson, 187 1 

1 87 1. Vesperus shiraziensis Dobson, J. Asiat. Soc. Bengal, ^o, 2: 459. Shiraz, 4,750 ft., 
South-Western Persia. 

Eptesicus serotinus andersoni Dobson, 1871 

1871. Vesperus andersoni Dobson, Proc. Asiat. Soc. Bengal, 211. Momein (Tengueh), 
Yunnan, China. Ranges to Fukien and Chekiang, Southern China. 

Eptesicus serotinus pallens Miller, 191 1 

191 1. Eptesicus serotinus pallens Miller, Proc. Biol. Soc. Washington, 24: 53. Cheng- 

yuanhsien, 70 miles west of Chingyangfu, Kansu, China. 
1929. Eptesicus serotinus pallidus Bobrinskii, Annu. Mus. Zool. Acad. St. Petersb. 

30: 235. 
Ranges to Shensi, Chihli, Shantung, in China; and Korea. 

Eptesicus serotinus meridionalis Dal Piaz, 1926 

1926. Eptesicus serotinus meridionalis Dal Piaz, Atti Soc. Ven. -Trent. Sci. Nat. 16: 63. 
Cagliari, Sardinia. 



Eptesicus serotinus intermedius Ognev, 1927 

1927. Eptesicus serotinus intermedius Ognev, J. Mamm. 8: 152. Murtasovo Station, 

near Madikavkaz, Terek region. Northern Claucasus. 

Eptesicus serotinus brachydigitus Mori, 1928 

1928. Eptesicus brachydigitus Mori, Zool. Mag. Tokyo, 40: 291 tin Japanese, 21 August 

1928). Annot. Zool. Jap. 2: 391 (in English, 20 December 1928). Heijo, 
Hcian, Nando, Korea. 
Tate also lists the form sinensis Peters (1880) as a race oi serotinus, but G. Allen 
placed it in the synonymy oi Nyctalus noctula plancei. 

Incerlae sedis 

Plptcsuus horikawai Kishida, 1924, Zool. Mag. Tokyo, 36: 127, 139. 

Eptesicus kohayashh Mori, 1928, Zool. Mag. Tokyo, 40: 292 (m Japanese, 21 
August 1928). Annot. Zool. Jap. 2: 392 (in English, 20 December). 
Heijo, Heian, Nando, Korea. 

Eptesicus aurijunctus (named as Vespertilw aurijimctus) Mori, 1928, Zool. Mag. Tokyo, 
40: 296 (in Japanese, 21 August 1928). Annot. Zool. Jap. 2: 393 (in Eng- 
lish, 20 December 1928). Keijo, Korea. ( Tuitalus aurijunctus Kishida & Mori, 
igs'i, Zool. Mag. Tokyo, 43: 372-391.) 

Eptesicus ranancnsis Kishida & Mori, Zool. Mag. Tokyo, 43: 379, nom. mid. Ranan, 
North Korea. 

Eptesicus tatei nom. nov. 

1863. Njcticeiits atratus Blyth, Cat. Mamm. Mus. Asiat. Soc. Bengal, 31. Eptesicus 
' atratus auct. but not atratus Kolenati, 1858, which = nilssonii. Darjeeling, 
North-Eastern India. 
This form is left incertac sedis by Tate (1942) and until the type is re-examined it is 
not possible to allocate the form with certainty. 

Genus NY CT ALUS Bowdich, 1825 

1825. Nrctalus Bowdich, E.xcursions in Madeira & Porto Santo, 36 (and footnote). 

Nyctalus verrucosus Bowdich. 
1829. Ptervgistes Kaup, Skizz. Europ. Thicrw. /.• 100. Vespertilw noctula Schreber. 
1842. Nocttdinia Gray, Ann. Mag. N.H. 10: 258. Contained two species, one of 

which is a synonym of V. noctula Schreber. 
1856. Paniigo Kolenati, AUg. Dtsch. Naturh. Ztg. 2: 131. Vcspertilio noctula Schreber 

and Vespertilio leisleri Kuhl. 

If this genus is considered congeneric with Pipistrellus, as by Simpson ( 1945), then 
Nyctalus has priority. 

The five species most likely to be valid in the area covered by this list arc: 

Nyctalus azoreum, page 159 .hyctulus leisleri, page 159 

.Hvctalu^ jojjrei, page 159 Myctaliis noctula, page 160 

Nyctaliis lasiopterus, page 160 

c;hiroptera — vespertilioninae 

Tate (1942, Bull. Amer. Mus. N.H. 80: 251) transferred N.joffrei to Pipistrellus, but, 
as remarked under that genus, we prefer tentatively to retain it in Nyctalus. Miller 
compared the other four species, all of which occur in Europe. Tate has shown that 
the prior name for the giant species is lasiopierus. Kuzyakin thought the form aviator 
was a valid species, but Tate makes it a subspecies of lasiopierus. From descriptions, 
the forms montanus and verrucosus seem very close to leisleri. Tate (1942, 256) states that 
the skull of montanus "exceeds considerably the measurements given by Miller for 
leisleri of Europe", but this seems an error; see Tate's table of measurements at the 
end of his paper. 

Nyctalus jojfrei group 
(Referred by Tate to Pipistrellus.) 

Nyctalus joffrei Thomas, 1915 

Approximate distribution of species: Burma. 

Nyctalus joffrei Thomas, 19 15 

1915. Nyctalus joffrei Thomas, Ann. Mag. N.H. /j.- 225. Kachin Hills, Upper Burma. 

Nyctalus noctula group 
( = restricted Nyctalus of Tate.) 

Nyctalus azoreum Thomas, 1901 
Appro.ximate distribution of species: Azores Islands, Atlantic. 

Nyctalus azoreum Thomas, 1901 

1901. Pterygistes azoreum Thomas, Ann. Mag. N.H. 8: 33. St. Michael, Azores 

Nyctalus leisleri Kuhl, 181 8 Lesser Noctule. Hairy-armed Bat 

Approximate distribution of species: England, Ireland, Holland, France, Switzer- 
land, Germany, Spain, Poland, Rumania; Russia, as far north as C. Volga and 
Moscow Province, and south to the Caucasus; Punjab, Kumaon. Perhaps repre- 
sented in Madeira by verrucosus. 

Nyctalus leisleri leisleri Kuhl, 1818 

1818. Vespertilio leisleri Kuhl, Ann. W^etterau Ges. Naturk. 4, i : 46. Hanau, Hessen- 

Nassau, Germany. 
i8i8. Vespertilio dasvkarpos Kuhl, loc. cit. 49, alternative name for leisleri. 
1839. yespertilio pachyonathus Michahelles, in Wagner, Schreber's Saugeth. Suppl. /, 

pi. 55b. Dalmatia. 

Nyctalus (?) leisleri montanus Barrett-Hamilton, 1906 

1906. Pterygistes montanus Barrett-Hamilton, Ann. Mag. N.H. ly: 99. Mussoorie, 
Kumaon, Northern India. 



Perhaps; the fulldwini^ form also represents hulen: 

Nyctalus verrucosus Bowdich, 1825 

1825. Nvctalus verrucosus Bowdich, Excursions in Madeira & Porto Santo, ':56 (and 

footnote). Island of Madeira. 
1 006. Nvctalus madcirae Barrett-Hamilton, Ann. Mae;. N.H. ly: C)8. Madeira. 

Nyctalus noctula Sthrebcr, 1774 Common Nortule 

Approximate distribution of species: Entjland, France, Switzerland, Spain, Italy, 
Norway, Sweden, Denmark, Holland, Germany, Rumania, Greece, Poland, Czecho- 
slovakia; Russia, southwards to the Caucasus, northwards to Leningrad district and 
Kirov (\'yatka) Province, Western Siberia, to the Altai and Tarbagatai Mountains, 
Usbekistan and Semircchyia, in Russian Turkestan; Kuldja, Western Chinese 
Turkestan; similar forms inhabit Chihli, Szcchuan, Fukien, and adjacent states in 
China; Japan; Nepal, Kashmir, Burma; Malay States; Persia, and Palestine accord- 
ing to Bodenheimcr. 

Nyctalus .\octula noctula Schreber, 1774 

1774. Vcspertiliu noctula Schreber, Saugeth. /.■ pi. 52 i text, p. 166). France. 

1776. Vcspcriilio lardarius Miiller, Natursyst. Suppl. Regist. Band, 15. France. 

1 781). Vespertilio magnus Berkenhout, Syn. Nat. Hist. Gt. Britain & Ireland, /.■ i. 
Cambridge, England. 

178c). ]'espfrtilio altivolaiu ^\'hite, X.H. & Antic|. of Sclborne, C)3. Selborne, Hamp- 
shire, England. 

(?) 1816. Vespertilio major Leach, Cat. Mamm. & Birds B.M. 5, nom. nud. 

1818. Vespertilio proterus Kuhl, Ann. W'etterau Ges. Naturk. 4, i: 41. Substitute for 

i82q. Vespertilio riijesceth Brehm, Isis, Jena, (]43. Jena, Thuringia, Germany. 

1844. Vespertilio palustris Crcspon, I'aune Mcridionale, /; 22. Marshes near Nimcs, 
Card, France. 

1869. I'esperugo noctula \ar. minima Fatio, Faune Vert. Suisse, /.■ 58. Geneva, Switzer- 

Range : Europe. 

Nyct.'^lus xoctul.v labl\tus Hodgson, 1835 

1835. Vespertilio labiata Hodgson, J. Asiat. Soc. Bengal, 4: 700. Nepal. Currently used 
for the form which occurs Kashmir, Darjeeling, Chin Hills (Burma) and, 
according to Chasen (1940), Malay States; but Tate (1942, 258) places it 
incertae sedis and states that he doubts whether it was based on a Nvctalus. 

Nyct.'^lus noctula plancei Gerbe, 1880 

1880. Vesperugo plancei Gerbe, Bull. Soc. Zool. France, 5.- 71. Pekin, Clhihli, China. 

1880. VcKperus sinensis Peters, Mber. Preuss. .\kad. Wiss. 258. Pekin, C^hiiia. 

Nyctalus noctul.\ princeps Ognev & Worobicv, 1923 

1923. Nyctalus noctula princeps Ognev & Worobiev, Fauna Vertebr. Mamm. Gvt. 
Moscou, 97. Voronej, Russia. 



Nyctalus noctula velutinus G. Allen, 1923 

1923. Nyctalus velutinus G. Allen, Amer. Mus. Nov. No. 85 : 7. Futsing, Fukien, China. 

Rane: China, states of Fukien, Chekiang, Kiangsu, Hupeh, Szechuan. 

Tate is inclined to treat plancei and velutinus as a valid, slightly smaller 

species than noctula. 

Nyctalus noctula meklenburzevi Kuzyakin, 1934 

1934. Nyctalus noctula meklenburzevi Kuzyakin, Bull. See. Nat. Moscou, 4^^: 323, 329. 
Tashkent, Russian Turkestan. 

Nyctalus noctula motoyoshii Kuroda, 1934 

1934. Nyctalus noctula motoyoshii Kuroda, in Siebold, Fauna Japonica (Japanese ed.), 

3: 3. {N.V.). Hondo, Japan. 
1934. Nyctalus noctula montanus Kishida, Lansania, Tokyo, 6, 52: 26. {N.V.). Not of 

Barrett-Hamilton, 1906. 

Nyctalus lasiopterus .Schreber, 1 780 Giant Noctule 

Approximate distribution of species: Switzerland, France (recorded 1932), Italy, 
Sicily; Russia, from Crimea and Transcaucasia as far north as Moscow Province 
and River Vetluga, east to Buzuluk Forest. The slightly smaller form, aviator, which 
Tate made a race, ranges widely in Japan and also occurs Shaweishan Island, off 
mouth of Yangtzekiang River, China (G. Allen). 

We follow Tate (1942) in the use of the name lasiopterus. Miller called it N.'maximus, 
and Ognev and Kuzyakin called it siculus; both are antedated by lasiopterus. 

Nyctalus lasiopterus lasiopterus Schreber, 1780 

1780. Vespertilio lasiopterus Schreber, in Zimmermann, Geogr. Gesch. 2: 412. No 

locality. ? Northern Italy (Chaworth-Musters). 
(?) 1827. Vespertilio ft.rugineus Brehm, Ornis, j: 26. Renthendorf, Thuringia, 


1868. Vespertilio noctula var. sicula Mina-Palumbo, Cat. Mammif della Sicilia. (N.V.) 


1869. Vesperugo noctula var. maxima Fatio, Faune Vert. Suisse, /.• 57. Amsteg, Uri, 

Range: Europe. 

Nyctalus (?) lasiopterus aviator Thomas, 191 1 

1840. Vespertilio molossus Temminck, Mon. Mamm. 2: 269. Not of Pallas, 1767. 

191 1. Nyctalus aviator Thomas, Ann. Mag. N.H. 8: 380. Tokyo, Hondo, Japan. 

Genus PIPISTRELLUS Kaup, 1829 

1829. Pipistrellus Kaup, Skizz. Europ. Thierw. /.• 98. Vespertilio pipistrellus Schreber. 
1838. Romicia Gray, Mag. Zool. Bot. 2: 495. Romicia calcarata Grav = Vespertilio kuhlii 




1856. Hypsugo Kolcnati, AUg. Dtsch. Xaturh. Ztg. 2: 131 [tnaunis ^ savii and 

kraschcnimkowii) . 
1856. Nannugo Kolcnati, loc. cit. Included W-spaiilw imthusii, V. kuhlii and ('. jnln- 

1867. Alohus Peters, Mbcr. Preuss. Akad. Wiss. 707. Vespcrtilio temnunckii Crctzsch- 

mar = Vespertilio ruppellii Fischer. Not of Leconte, 1856. 
1875. Scotozous Dobson, P.Z.S. 372. Scotozous dormeri Dobson. Valid as a subgenus. 
1899. Euvesperugo Acloque, Faune de France, Mamm. 35 (part). (Included si.\ 

species, one of which was I', pipistrellus.) 
iqo2. la Thomas, Ann. Mag. N.H. 10: 163. la io Thomas. \'alid as a subgenus. 
191 7. Megapipiilrellus Bianchi, Annu. Mas. Zool. St. Petersb. 21: Ixwii. Pipistrellus 

annectcns Dobson. \'alid as a subgenus. 
1926. Eplencops Roberts, Ann. Transvaal Mus. //.• 245. Scotophilus rusluus Tomes, 

from South-West Africa. Valid as a subgenus. 
1946. Vansonia Roberts, Ann. Transvaal Mus. 20: 304. Pipistrellus rernavi Roberts, 

from Bechuanaland. (A race of rUppellii, fide G. Allen.) 

Kuz\akin, in Bobrinskii, 1944, refers this genus, and I-lptesicus, to the earlier-named 
genus Vespertilio, stating that it is a large and extremely heterogeneous group but that 
the features of its individual representatives are so closely interlocked that it is not 
practicable to divide the group into independent genera as previous writers have 
done. The main difficulty seems to be that in the U.S.S.R., the small upper premolar, 
characteristic of Pipistrellus, may be absent in Pipistrellus savii as understood by 
Kuzyakin. Kuzyakin states that a number of forms have been described, classified as 
"species" or even "genera" (Vesperugo caucasicus, Amblvotus tauricus, A. velox, Epiesicus 
alaschanicus, etc.), b>it they have all proved to be simply types of individual and 
geographical variation in one species. He recognizes three forms in the U.S.S.R., 
P. s. savii, always with an upper small premolar tooth; P. s. alaschanicus, "half of the 
individuals have small upper premolar teeth and half do not", and P. s. caucasicus, 
"small upper premolar is missing in nearly all cases". In P. savii (Russian races) the 
penis is bent iiit" the shape of an inverted L ("unlike all other Vespertilio as understood 
bv Kuzyakin) ; this is an alternati\e character given by this author to divide savii 
from other species, whether individually it has the small upper premolar or not. 
Strictly speaking, Pipistrellus is not more than a subgenus of Eptesicus, which itself 
mitrht well be referred to Vespertilio. But whereas in Russia the suppression of these 
two convenience genera does not make much difference (only about a dozen species 
are involved), when the problem is looked at from a world point of view it becomes 
more difficult. For instance, Pipistrellus is such a major division in the Old World 
tropics that Tate, in his review of the Vespcrtilionidae, makes it typify an entire 
generic assemblage ("Pipistrelli"). We do not feel that American authors, or students 
of Africa and the Indomalayan region, would take a very good view < if lumping such 
a large number of spciics into Vespertilio. Therefore, for convenience only, and bear- 
ing in mind that an alt(rnati\e character is gixcn wliich will separate Pipistrellus savii 
in the U.S.S.R. hum fithcr Rus-^ian bats, we list Pipi\lrelhn and Eptesicus. follnwing 
Miller and Tate. 

Hollistcr has pointed out that the characters used b\ Miller for the genus Scotnzous 




are not of generic value. The name is currently placed in synonymy, although Tate 
(1942) retains it for the Indian species, dormcri. Surely it is at most a subgenus, and 
the same applies to la, as already indicated by Simpson, and suggested by Tate 
(P- 259)- 

Tate recognizes and defines 12 species groups of the present genus in the Palae- 
arctic and Indian region, and two more, typified by the Indian Scotozous and la, arc 
here added. 

The 21 species most likely to be valid in the present region are: 

Pipistrellus abramus, page 165 
Pipistrellus affinis, page 167 
Pipistrellus anneckns, page 1 72 
Pipistrellus ariel, page 1 7 1 
Pipistrellus babu, page i6g 
Pipistrellus ceylonicus, page 167 
Pipistrellus circumdatus, page 1 7 1 
Pipistrellus coromandra, page 165 
Pipistrellus deserti, page 169 
Pipistrellus dormeri, page 172 
Pipistrellus io, page 173 

Pipistrellus kuhli, page 168 
Pipistrellus lophurus, page 167 
Pipistrellus maderensis, page 1 7 1 
Pipistrellus mimus, page 1 66 
Pipistrellus mordax, page 1 7 1 
Pipistrellus nathusii, page 1 64 
Pipistrellus pipistrellus, page 1 63 
Pipistrellus pulveratus, page 167 
Pipistrellus riippelli, page 172 
Pipistrellus savii, page 169 

According to Tate, Pipistrellus tralntitius Horsfield ' 1824, Vesperugo tralatitius Hors- 
field, Zool- Res. Java, from Java), which was recorded from Tonkin, Indo-China, by 
Osgood, was based on a Myotis. 

Tate transfers stenopterus and joffrei from Nyctalus to Pipistrellus, but we do not feel 
inclined to follow him in this classification, for two reasons: Miller (1907) definitely 
placed stenopterus (from Borneo) in Nyctalus, stating that he had examined "all the 
known species", and Thomas, in describing jo/r« stated that the proportions of the 
digits were as in Nyctalus [Nyctalus differing from Pipistrellus chiefly in its shortened 
fifth finger). 

Authors who wish to merge Pipistrellus with Nyctalus should note that Nyctalus takes 

Subgenus PIPISTRELLUS Kaup, 1829 
Pipistrellus pipistrellus group 

Pipistrellus pipistrellus Schreber, 1774 Common Pipistrelle 

Approximate distribution of species: Britain, Ireland, Sweden, Denmark, \orway, 
France, Switzerland, Italy, Spain, Sardinia, Germany, Holland, Rumania, Yugo- 
slavia, Poland, Greece; Russia, from the Caucasus, north roughly to the levef of 
Moscow; Russian Turkestan, where widely distributed. Has been recorded from 
Japan, Formosa and Korea (Kuroda). Asia Minor (B.M.), Persia; Kashmir. Recorded 
from Morocco (1933). 



1774. I'fspfililio pipislifHiis Schreber, Siiugeth. /, pi. 54 (text, p. 167). France. 
1776. Vespertilio pipistrelle Muller, Natursyst. Suppl. Regist. Band, 16. 
i8-;3. I'fspfiiilio prgmaeiis, Zool. J. /.■ 559. Dartmoor, Devonshire, England. 
1834. Ves/Jfi'lilio hiachyotoi Baillon, Mem. Soc. £mul. Abbeville, /(JjJjJ." 50. Abbeville, 

Somme, France. 
(?) 1838. Scotophilus murinus Gray, Mag. Zool. Bot. 2: 497. 
(?) 1839. VespcrtiUo {Ptpistrcllus) pipislrellus var. nigra de Sclys Longchamps, fitudes 

de Slicromamm, 140, nom. mid. 
(?) 1839. Vesperlilw [Pipislrellus) pipistrellus var. rufescens de Selys Longchamps, loc. cit. 

ngm. nud. Not of Brehm, 1829. 
1840. VespcrtiUo piisilliis Schinz, Fauna Europ. /.■ 9. 

1840. Vespertilin melanopterus Schinz, loe. cit. Rcnthendorf, Thuringia, Germany. 
1840. Vcsperlilin stenotus Schinz, loe. cit., same locality. 
1840. Vespcrtilio minutissimus Schinz, loc. cit. Zurich, .Switzerland. 
I?) 1842. Kerivoula gnseus Gray, Ann. Mag. .\.H. ifi: 258. No locality. 
184',. Pipistrellus nigricans Bonaparte, Atti della scsta Riun. dcgli Sci. Italiani, Milano, 

1844: 340. Sardinia. 
1843. Pipistrellus genei Bonaparte, loc. cit., alternative for nigricans. 
184'"^. Pipislrellus trpus Bonaparte, loc cit., substitute ior pipistrellus. 

1862. Vesperugo pipistrellus xvlw uiacropterus ]c\Uc\c^, X'c'h. Zool. Bot. Ges. \\'icn, 12: 

250. Kaschau, Hungary. 

1863. J^'annugo pipistrellus var. flarescens Koch, Jb. Nassau \'er. Naturk. 18: 41)1. 

Nassau, Germany. 
1863. Nannugo pipistrellus var. nigricans Koch, loc. cit., not of Bonaparte, 1845. Nassau, 

1863. J^annugo pipistrellus \ar. limbatus Koch, loc. cit. 492. Siegen, Nassau, Germany. 

1904. Pipistrellus pipistrellus mediterrancus Cabrera, Mem. Soc. Esp. H.N. 2: 273. 

Valencia, Spain. (Placed in synonymy by Miller, but regarded by Tate 
(1942) as a race oi nathusii.) 
Range; Europe, Asia Minor, Persia. 

Pipistrellus pipistrellus b.^iCtrianus Satunin, 1905 

i?) 1840. Vespertilio lacteus Temminck, Mon. Mamm. 2: 245. Locality unknown. 
1873. Vesperugo akokomuli var. almatensis Severtzox', Mem. Soc. Amis. Sci. Nat. 

Moscou, 8, 2: 79; 1876. Ann. Mag. N.H. 18: 42, nom. nud. Turkestan. 
1882. Vespertilio oxianus Bogdanov, Outline N.H. khibinskoyo, 78, nom. nud. (M.V.) 

1905. Pipistrellus hactrianus Satunin, Mitt. Kaukas. Mus. 2: 67, 85. Oasis of Tedzen, 

Transcaspia, Russian Turkestan. 
Range includes Gilgit, Kashmir. 

Pipistrellus nathusii Kcyserling & Blasius, 1839 Nathusius' Pipistrelle 

Approximate distribution of species: France, Switzerland, Spain, Italy, Germany, 
Poland, Holland, Denmark, Hungary; Russia, from Leningrad and lower Vyatka 
River, east to Orenburg, south to Black Sea and Caucasus. ? Persia; Palestine 
according to B<idenhcimer. 



PiPisTRELLUS NATHUsii Kcyscrling & Blasius, 1839 

1839. V^spertilio nathusii Keyserling & Blasius, Arch. Naturgesch. 5, i : 320. Berlin, 

1905. Vesperugo nathusii var. unicolor Fatio, Arch. Sci. Nat. Geneve, ig: 510. Geneva, 

Pipistrellus abramus group 

Pipistrellus abramus Tcmminck, 1840 Japanese Pipistrelle 

Approximate distribution of species: Southern Ussuri region, Eastern Siberia; 
Japan; China, states of Chihli, Shantung, Fukien, Szechuan, Hupeh, Hunan; 
Hainan, Indo-China; Formosa; Java, Banka (see Tate, 1942, 237). Probably the 
Burmese form below may be regarded as a race. 

Pipistrellus abramus abramus Temminck, 1840 

1840. Vespertilio abramus Temminck, Mon. Mamm. 2: 232, pi. 58, figs, i, 2. Nagasaki, 

Kiushiu, Japan. 
(?) 1840. Vespertilio akokornuli Temminck, Mon. Mamm. 2: 233, pi. 57, figs. 8, 9. 

1842. Vespertilio irretitus Cantor, Ann. Mag. N.H. g: 481. Chusan Island, Chekiang, 

1857. Seotophilus pumiloides Tomes, P.Z.S. 51. China. 
1922. Scotophilus pomiloides Mell, Arch. Naturgesch. 88a, 10: 14. 
Range: Japan and China, as above; Annam, in Indo-China. 

Pipistrellus (?) abramus paterculus Thomas, 1915 

1915. Pipistrellus paterculus Thomas, J. Bombay N.H. Soc. 2^: 32. Mt. Popa, Upper 
Burma. Range includes Shan States and Chindwin, Burma. 

Tate also refers the following named form to the present group : 
Pipistrellus camortae Miller, 1902 

1902. Pipistrellus camortae Miller, Proc. U.S. Nat. Mus. 2^: 779. Kamorta Island 
Nicobar Islands, Bay of Bengal. 

Pipistrellus coromandra group 

It is not impossible that the name P. imbricatus Horsfield, 1824, Java, is the prior 
name in this group. 

Pipistrellus coromandra Gray, 1838 Indian Pipistrelle 

Approximate distribution of species: Fukien, in Southern China, Hainan; Indo- 
China; Burma, Bhutan Duars, Sikkim, Kumaon, many localities in Peninsula of 
India, and Ceylon. Persia, \i aladdin is rightly allocated here. 



1838. Scotophilus coromandra Gray, Mag. Zool. Bot. 2: 498. Pondichcrry, Coromandel 
coast, India. 

1851. Vespertilio coromandelicus Blyth, J. Asiat. Soc. Bengal, 20: 159. 

1853. Mvotis parvipes Blyth, J. Asiat. Soc. Bengal, 22: 581. Masori (? Mussoorie, 

(?) 1855. Vespcrugo blythii Wagner, Schreb. Saugcth. Suppl. 5.- 742. Ceylon. 

1863. Scotophilus coromandelianus Blyth, Cat. Mamm. Mus. Asiat. Soc. Bengal, 33. 

(?) 1872. Vesperugo micropus Peters, P.Z.S. 708. Dchra Dun, near Simla, North- 
Western India. Tate lists this form as possibly valid. 

Range: Cevlon, north to Kumaon and Bhutan Duars. 


1905. /'!/;/.(/;-(7/«_s fl/afl'(/w Thomas, Abstr. P.Z.S. No. 24: 23. 1906, P.Z.S. /505, 2:521. 

Derbent, 30 miles west of Isfahan, 6,500 ft., Persia. 


1906. Pipistrellus porlaisis ]. Allen, Bull. Amcr. Mus. N.H. 22: 487. Portcn, Island of 



1928. Pipiitrclliis coromandrus tramatus Thomas, P.Z.S. 144 Thai-nien, Tonkin, Indo- 
China. Range includes Annam, Laos, and Fukien in Southern China. Tate 
suggests it may be the same as portnnii. Anthony ( 1941) recorded it from 
Northern Burma. 

Pipistrellus tenuis group 
Based on P. tenuis Temminck, 1840, from Java (e.xtralimital). 

Pipistrellus mimus Wroughton, 1899 Indian Pygmy Pipistrelle 

Approximate distribution of species: Ceylon, most of Peninsular India, Kathiawar, 
Palanpur, Cutch, Sind, Punjab, Sikkim, Bhutan Duars, Assam, Burma; .^nnam, 


1899. Pipistrellus mimus Wroughton, J. Bombay N.H. Soc. 12: 722. Mheskatri, 
Dangs, Sural district. Western India. Range: south to Ceylon, north to 
Kathiawar and district, Kumaon, Sikkim, east to Western Burma and 

PiPISTRELLUS MIMUS oi.AUCiLLUS ^\'rou<;;hton, 1912 

1912. PifnUrelhn mimus glaucillus Wroughton, J. Bombay N.H. .Soc. 21: ]6(). Multan, 
Punjab, India. Ranges to Sind. 


I(|I5. Pi/mlrellin /irtneifnilus Thomas, ,\nn. .\Ia^. N.H. ij: 231. Gauhati, .\ssam. 


Pipistrellus affinis group 

Pipistrellus affinis Dobson, 1871 Chocolate Bat 

Approximate distribution of species: Bhamo (Yunnan-Burma border). 

Pipistrellus affinis Dobson, 1871 

1 87 1. Vesperugo (Pipistrellus) affinis Dobson, Proc. Asiat. Soc. Bengal, 213. Bhamo, 
North-Eastern Burma. Tate also records it from the Likiang Range, Yun- 
nan, China. 

Pipistrellus pulveratus Peters, 1871 

Approximate distribution of species: Szechuan, Yunnan and Fukien, in Southern 

Pipistrellus pulveratus Peters, 1871 

1871. Vesperugo pulveratus Peters, in Swinhoe, P.Z.S. i8yo: 618. Amoy, Fukien, China. 

Pipistrellus lophurus Thomas, 1915 

Approximate distribution of species: Tenasserim. 

Pipistrellus lophurus Thomas, 19 15 

191 5. Pipistrellus lophurus Thomas, J. Bombay N.H. Soc. 23: 413. Maliwun, Victoria 
Province, Tenasserim. 

Pipistrellus cerlonicus group 

Pipistrellus ceylonicus Kelaart, 1852 Kelaart's Pipistrelle 

Approximate distribution of species: Ceylon, Peninsula of India (where widely 
distributed), north to Kathiawar, Sind, Cutch, Bengal. Probably represented in 
Burma and Indo-China. 

Pipistrellus ceylonicus ceylonicus Kelaart, 1852 

1852. Scotophilus ceylonicus Kelaart, Prodr. Faun. Zeylan, 22. Trincomalee, Ceylon. 

Pipistrellus ceylonicus indicus Dobson, 1878 

1878. Vesperugo indicus Dobson, Cat. Chiroptera B.M. 222. Mangalore, Malabar 
coast, India. Range: Southern Peninsular India. 

Pipistrellus ceylonicus chrysothrix Wroughton, i8gg 

1899. Pipistrellus chrysothrix Wroughton, J. Bombay N.H. Soc. 12: 720. Mheskatri, 

Surat Dangs, India. Range: northwards from the range of indicus, south of 

that of subcanus, to Bengal. 

Pipistrellus (?) ceylonicus raptor Thomas, 1904 

1904. Pipistrellus raptor Thom3.s, Ann. Mag. N.H. 13: 387. Tonkin, Indo-China. 


PALA1:ARC:TIC and IXDIAX mammals 1738 1946 

1915. Pipislrelliis shanorum Thomas, J. Bombay N.H. Soc. 24: 29. Pyaunsgauiic;, 
Northmi Shan States, Burma. 


1 915. Pipistrdhis cnlonicus suhcanus Thomas, J. Bombay N.H. Soc. 24: 30. Wilala, 
Junaa;adh, Kathiawar, India. Range inckides Sind, C'utch, Palanpur. 

PipistrcUiis kiihli group 

Pipistrellus kuhli Kuhl, 1819 Kuhl's Pipistrellc 

Approximate distribution of species: France, Germany, Switzerland, Italy, Spain, 
Balearic Islands, Sardinia, Greece; Crimea, Caucasus and lower Amu-Darya, in 
Russian Turkestan; Asia Minor (B.M.), Persia, Afghanistan, Palestine, Arabia; Sind, 
Kashmir; Egypt, Algeria, Morocco. Also known from Asben, Ken\a and South 
Africa to Transvaal, Natal and Knysna, Cape Province. 

Pipistrellus kuhli kuhli Kuhl, 1819 

1819. Vesperlilio kuhlii Kuhl, Ann. W'cttcrau. Ges. Naturk. 4, 2: 199. Trieste (Italian- 
Yugoslavian border). 

1829. Vespertilio pipistniliis var. aegrpliiis Fischer, Synops. Mamm. 105. Thebes, 


1830. Vcipertilw marginatus C'retzschmar in Ruppcll. Atlas Rcisc nordl. Afrika, 

Saugcth. 74, pi. 29a. "Arabia Pctraca" (Sinai) and Nubia, Sudan. According 
to Anderson & de Winton, 1902, Zool. Egypt, Mamm. 127, from Egypt. 

1835. Vespertilio albolimhatus Kiister, Isis, Jena, 75. Cagliari, Sardinia. 

1837. Vespertilio vispistrellus Bonaparte, Faun. Ital, /, fasc. 20. Sicily. 

1837. Vespertilio alcrthoe Bonaparte, loc. eit. fasc. 21. Sicily. See Miller, 1912, 215. 

1838. Romicia calearata Gray, Mag. Zool. Bot. 2: 495. Locality unknown. 

1840. Vespertilio ursula Wagner, Schreb. Saugeth. Suppl. /.• 505. Morea, Greece. 

1841. Pipistrellus marginatus Bonaparte, Faun. Ital., Indie, distrih. Substitute for 

1863. .^'vctieelus caniis Blyth, Cat. Mamm. Mus. Asiat. Soc. Bengal, 32. India. Tate 

suggests this may be a valid race. 
1867. Pipistrella minuta Lochc, Expl. Sci. Algerie, Zoo]., Mamm. 78. Oasis of Messad, 

Southern Algeria. 
(?) 1867. Seotophilus lobatiis jcrdon, Mamm. Ind. 35. Madras, India. 
1872. Vespertilio {Pipistrellus) leucotis Dobson, J. Asiat. Soc. Bengal, 41: 222. Rajanpur, 

Punjab, North-Western India. 
1886. Vespertilio kuhlii var. albicans Monticelli, Atti Soc. Ital. Sci. Nat. 2j: 200. 

Caivano, Naples, Italy. 
1886. Vespertilio kuhlii var. pullatus Monticelli, loc. cil. Bella Vista, near Portici, Naples, 

Range: Europe, North Africa, Sind, Persia. 

Pipistrellus kuhli lepidus Blyth, 1845 

1845. Pipistrellus lepidus Blyth, J. Asiat. Soc. Bengal, 14: 340. Kandahar, Afghanistan. 
Ranges to Kashmir and Upper Sind frontier. 



PipiSTRELLUs KUHLi iKHWANius CheesiTian & Hinton, 1924 

1924. Pipistrellus kuhlii ikhwanius Cheesman & Hinton, Ann. Mag. N.H. 14: 549. 
Hufuf, Hasa, Central Arabia. 

Pipistrellus kuhli pallidus Heim de Balsac, T936 

1936. Pipistrellus kuhlii pallidus Heim de Balsac, Bull. Biol. Paris, 21, SuppL: 180. 
Northern Sahara to the A'haggar, Algeria. 

Pipistrellus babu Thomas, 19 15 

Approximate distribution 'of species: Punjab, Kumaon, Nepal, Sikkim, Bhutan 
Duars, Assam and Central Provinces, India. 

This species is included provisionally in the kuhli group by Tate. It differs in having 
a long outer incisor, and has P 2 not so strongly displaced internally. 

Pipistrellus babu Thomas, 1915 

1915. Pipistrellus babu Thomas, J. Bombay N.H. Soc. 24: 30. Murree, 8,000 ft., 

Pipistrellus deserti Thomas, 1902 

Approximate distribution of species: Libya. 

In describing this species, Thomas compared it with P. kuhli, from which it differed 
in its shorter skull and toothrow, and narrower braincase. Miller (1907) placed it in 
Scotozous, but Thomas & Hinton, 1923, P.^-S. 250, confirmed Thomas's earlier 
opinion that it was allied to kuhli. Dentition as in P. kuhli, but size smaller. 

Pipistrellus deserti Thomas, 1902 

1902. Pipistrellus deserti Thomas, P.Z.S. igo2, 2: 4. Mursuk, Tripoli, Libya. 

Pipistrellus savii group 

Pipistrellus savii Bonaparte, 1837 Savi's Pipisti'elle 

Approximate distribution of species: France, Switzerland, Italy, Spain, has been 

recorded from Germany (Breslau); Greece; Crimea, Caucasus, Turkestan (Turk- 

menia, Ust-Urt, Tianshan, etc.), and Ussuri region of Eastern Siberia; Mongolia; 

Sikkim, Assam, Burma (but Tate doubts whether the India named forms really 

represent the species) ; Canary Islands. 

Kuzyakin states that the forms caucasicus, tauricus, velox and alaschanicus, hitherto 

regarded as small members of Eptesicus, represent this species. 

Pipistrellus savii savii Bonaparte, 1837 

1837. Vespertilio savii Bonaparte, Faun. Ital. /.• fasc. 20. Pisa, Italy. 

1837. Vespertilio aristippe Bonaparte, loc. cit., fasc. 21. Sicily. 

1837. Vespertilio leucippe Bonaparte, loc. cit. Sicily. 

1838. Vespertilio bonapartii Savi, Nuovo Giorn. Lett. Pisa, ^7.- 226. Pugnano, near 

Pisa, Italy. 




1844. Vespertilio nigrans Crespon, Faunc Mcridionale, /; 24. Nimes, Gard, France. 
1853. Vespirlilio maurus Blasius, Arch. Naturgesch. /g, 1 : -s,-^. Central chain of Alps. 
1872. Vfsperlilio a«ilis Fatio, Faune Vert. Suisse, /.• appendix, iii. New name for savii. 
1004. Vespertilio ochromixtus Cabrera, Mem. Soc. Esp. H.N. :?.• 267, pi. 3, figs, i & 4. 

Sierra de Guadarrama, Madrid, Spain. 
Range; Europe. 


1859. Scolnphili/s )l(irwini Ton^Ci, P.Z.S. 70. Las Palmas, Canary Islands. A\ailable if 
the Clanary Islands form should prove distinct. 


1 87 1. Pipistrdlus austcnianus Dobson, Proc. Asiat. Soc. Bengal, 213. Cherrapunjee, 
Khasi Hills, Assam. Range: to Shan States, Burma. 

PiPisTRELLUs (.'') .s.wii c.\ucAsicus Satuuiu, 1901 

1901. Vespeniuo [Vcspcrus) cnucaucus Satunin, Zool. Anz 24: 462. Tiflis, Caucasus. In 

placing this form here we follow Kuzyakin, in Bobrinskii (1944, loi). 

Range: tn C'rimea and Turkestan. 


Kjib. Pipistnllus cadornae Thomas, J. Bombay N.H. Soc. 24: 416. Pashok, 3,500 ft., 
Darjeeling. North-Eastern India. 

PiPISTRELLUS (?) S.'VVII TAMERLAM Bobrillskii, I918 

1918. Epicsicus tamerhvn Bobrinskii, Fauna & Flora P.ussia, ifj: 13-16. 'N.V.) 
Baisunski Bay, Bcjkhara district, Russian Turkestan. Placed (as a race) in 
E. (aiicasuiii by Ognev, which is included in P. savii by Kuzyakin ! 1944). 


192G. Eptciicus caiicauciii pallescens Bobrinskii, C.R. Acad. Sci. U.R.S.S., A, 97. River 
Moldja, northern slope of Kotan Tagh, Southern Sinkiang. Des- 
cribed as a race o{ caucasicus which Kuz)akin (1944) refers to the present 

PiPLSTRELLUS (?) S.A.VII ALASCHANicus Bobrinskii, 1926 

1926. Eptcsicus alaschanicus Bobrinskii, C.R. Acad. Sci. U.R.S.S., A, 98. Pass of 

Hotin Gol, near Dinyuanin, western slope of Alashan Range, Mongolia. 
Ranges to Ussuri district. Eastern Siberia. Kuzyakin (1944) lists this as a 
\alid race of P. saini. 


1927. Awhlrnliis lauricus Ognev, j. Mamm. 8: 153. Karadagh, Crimea. Referred to 

the present species by Kuzyakin (1944), but not regarded as a valid race. 
Perhaps —: cauc/niai^. 



1927. Amblyotus velox Ognev, J. Mamm. 8: 154. Vladivostock, Eastern Siberia. 
Referred to savii by Kuzyakin (1944) but not regarded as a valid race. 
Perhaps = alaschanicus. 

The following two African species are mentioned by Tate in the present group; 
both are likely to be valid. P. maderensis was compared with savii by Dobson. P. ariel 
(a pygmy species, forearm 30 mm., total length of skull 11. 3 mm.) differs from P. 
deserti apparently in narrower braincase and shorter toothrow; its outer upper incisor 
is unusually long, and it has P 2 extremely reduced, as in P. savii. 

Pipistrellus maderensis Dobson, 1878 

Approximate distribution of species: Madeira and Canary Islands. 

Pipistrellus maderensis Dobson, 1878 

1878. Vesperugo maderensis Dobson, Cat. Chiroptera B.M. 231, pi. 12, fig. 5. Island of 

Pipistrellus ariel Thomas, 1904 

Approximate distribution of species: Southern Egypt. 

Pipistrellus ariel Thomas, 1904 

1904. Pipistrellus ariel Thomas, Ann. Mag. N.H. 14: 157. Eastern Egyptian Desert, 
22° N., 35° E., 2,000 ft. 

Pipistrellus circumdatus group 

Pipistrellus circumdatus Temminck, 1840 Large Black Pipistrelle 

Approximate distribution of species: Java; Northern Burma (Anthony, 1941) and 
" India " (Dobson, Blanford). 

Pipistrellus circumdatus Temminck, 1840 

1840. Vespertilio circumdatus Temminck, Mon. Mamm. 2: 214. Tapos, Java. 

Pipistrellus mordax Peters, 1866 

Approximate distribution ofspecies: Java; Kumaon, Darjeeling, Calcutta, Ceylon. 

Pipistrellus mordax Peters, 1866 

(?) 1843. Scotophilus maderaspatanus Gray, List Mamm. Coll. B.M. 29, noin. nud. 

Madras, India. 
1866. Vesperugo mordax Peters, Mber. Preuss. Akad. ^Viss. 402. Java. 

M 171 

Subgenus MEGAPIPISTRELLUS Bianchi, 1916 

Pipistrellus annectens Dobson, 1871 Intermediate Bat 

Approximate distribution of species: Assam, and has been recorded from Sumatra. 

Pipistrellus annectens Dobson, 1871 

1 87 1. Pipistrellus annectens Dobson, Proc. Asiat. Soc. Bengal, 213. Naga Hills, Assam. 

1876. Wsperugo annectens Dobson, Monogr. Asiat. Chiroptera, 116. 

Subgenus SCOTOZOUS Dobson, 1875 

Tate referred only dormcri here, and treated the riippelli group, as a group of Pipi- 
strellus. For note on the dental characters of the two species, see Miller, 1907, Families 
& Genera of Bats, 206. 

Pipistrellus riippelli group 

Pipistrellus riippelli Fischer, 1829 Ruppell's Bat 

Approximate distribution of species: Sudan, Uganda, Angola, Bcchuanaland; 
north to Egypt; Iraq. 

Pipistrellus rlippelli ruppelli Fischer, 1829 

1826. Vespertilio temminckii Cretzschmar, in Ruppell, Atlas Reise. nordl. Afrika, 

Siiugeth. 17, pi. 6. Not of Horsfield, 1824. 
1829. Vespertilio ritppellii Fischer, Synops. Mamm. log. Dongola, Anglo-Egyptian 

Ranges north to Eg^'pt. 

Pipistrellus ruppelli coxi Thomas, 1919 

191 9. Pipistrellus coxi Thomas, J. Bombay N.H. Soc. 26: 747. Beit Mahommad, 
Amara, Irac|. 

Pipistrellus dormcri group 

Pipistrellus dormeri Dobson, 1875 Dormer's Bat 

Approximate distribution of species: India, from Cutch, Kathiawar, Bengal, 
Bhutan Duars, south to Bombay, Dharwar and Bellary in the Peninsula; Formosa 

Pipistrellus dormeri dormeri Dobson, 1875 

1875. Scotozous dormeri Dobson, P.Z.S. 373. Bellary Hills, India. Range: as in the 
species, except Kathiawar, Cutch, Palanpur. 


191 5. Scotozous dormeri caurinus Thomas, J. Bombay X.H. Soc. 24: 33. Junagadh, 
Kathiawar, 400 ft., India. Ranges to C\Uch, Palanpur. 


Subgenus lA Thomas, igo2 

Pipistrellus io Thomas, 1902 Great Pipistrelle 

Approximate distribution of species: Szechuan, Hupeh and Kweichow, China. 

Pipistrellus 10 Thomas, 1902 

1902. la 10 Thomas, Ann. Mag. N.H. 10: 164. Chungyang, Southern Hupeh, China. 

Another named species o{ Pipistrellus is P. anthonyi Tate, 1942, which is placed by 
him in the so-called "Pipistrellus joffrei group". Its status seems not absolutely clear. 
The species jq^m has hitherto been regarded as a Nyctalus, together with the Bornean 
N. stenopterus which Tate also refers to the "P. joffrei group". As noted above, we prefer 
for the present to leave J^'. joffrei in the genus Nyctalus. The proportions of the digits 
are not stated in the original description of P. anthonyi. 

Pipistrellus anthonyi Tate, 1942 

1 94 1. Pipistrellus affinis Anthony , Field Mus. Publ. Zool. zy: 81. Not ofDobson, 1871. 

1942. Pipistrellus anthonyi Tate, Bull. Amer. Mus. N.H. 80: 252. Changyinku, 

7,000 ft.. Northern Burma. 

Genus GLISCHROPUS Dobson, 1875 
1875. Glischropus Dobson, P.Z.S. 472. Vesperugo tylopus Dobson. 

1 species in the area covered by this list: 

Glischropus tylopus, page 173 

Simpson ( 1 945) suggests that this should be included in Pipistrellus, and it may well 
be only a subgenus of that. It is, according to Tate, "an offshoot o^ Pipistrellus in 
which the apparatus for grasping has undergone modification". 

Glischropus tylopus Dobson, 1875 Thick-thumbed Pipistrelle 

Approximate distribution of species: Burma; Malay States, Sumatra, Borneo, 
perhaps to Philippine Islands. 

Glischropus tylopus Dobson, 1875 

1875. Vesperugo tylopus Dobson, P.Z.S. 473. North Borneo. Ranges north to Karen 
Hills, Eastern Lower Burma. 

Genus HESPEROPTENUS Peters, 1868 

1868. Hesperoptenus Peters, Mber. Preuss. Akad. Wiss, 626. Vesperus doriae Peters, 
from Borneo. 

2 species in the area covered by this list: 

Hesperoptenus blanfordi, page 174 
Hesperoptenus tickelli, page 174 



These U\'0 species differ conspicuously from each other in size, tickcUi bcinj; much 
the larger. 

The genus as understood by Miller ( 1907) is characterized by the peculiar position 
of the second upper incisor, a character which is said to be present in both the Indian 
species. But Miller, and subsecjuent authors, do not seem to have examined the type 
species, and it may be that this genus will pro\e untenable in the sense in which it is 
at present accepted. Tate (1942) notes that hlanjordi has a digital adaptation similar 
to that of Glischrojiiis. 

Hesperoptenus tickelli Blyth, 1851 Tickelfs Bat 

Approximate distribution of species: India — Rajputana, Orissa, Bombay, Madras, 
Ceylon, Bengal, Bhutan Duars. (Blanford also quoted it from the Andaman Islands 
and Moulmein district, Burma.) 

Hesperoptenus tickelli Blyth, 1851 

1851. .\vclicejus tickelli Blyth, J. Asiat. Soc. Bengal, 20: 157. Chaibassa, Orissa, India 

(Wroughton, 19 10). 
(?) 1851. Nvcticejus isabelUniis Horsfield, Cat. Mamm. Mus. E. Ind. Co. 38. No 


Hesperoptenus blanfordi Dobson, 1877 Blanford's Bat 

.\pproximate distribution of species: Tenasscrim, Malay Peninsula. 

Hesperoptenus bl.'\nfordi Dobson, 1877 

1877. Vesperugo {Hesperoptenus) blanfordi Dobson, J. Asiat. Soc. Bengal, 4G, 2: 312. 

Genus TYLONYCTERIS Peters, 1872 

1872. Trlonvcleris Peters, Mber. Preuss. Akad. Wiss. 703. Vespertilio pachjpiis Tem- 

2 species in the area covered by this list: 

Trlonvcterii paehrpu^, page 174 
Trlonvclcrn rnbuslula, page 1 75 

Review: Tate, 1942, £;///. Amrr. Mus. JS'.H. 80: 266, wherein two groups of species 
(a larger and a smaller) arc shown to occur together. 

Tylonycteris pachypus Temminck, 1840 Club-footed Bat 

Approximate distribution of species: Yunnan and ? Kwantung, in Southern China, 
Burma, Manipur, Sikkim; Dharwar, southwards to Coorg in South- Western India; 
Tonkin, Laos and Annam, in Indo-China, Malay States, Borneo, Java, Bali, 
Sumatra fTate), to Luzon. Philippine Islands. : Blanford also quoted it from the 
.■\ndaman Islands.) 



(Tylonycteris pachypus pachypus Temminck, 1840. Extralimital) 
1840. Vespertilio pachypus Temminck, Mon. Mamm. 2: 217, pi. 54, figs. 4-5-6. 
Bantam, Western Java. 

Tylonycteris pachypus fulvid.a. Blyth, 1859 

1859. Scotophilus fulvidus Blyth, J. Asiat. Soc. Bengal, 28: 293. Schwegyin, Sittang 

River, South-Eastern Burma. 
1915. Tylonycteris rubidus Thomas, Ann. Mag. N.H. 75.- 227 (error hr fulvidus). 
Range: Sikkim, Manipur, Chin Hills, Shan States, Pegu, Tenasserim, Yunnan, Laos, 
Tonkin, Annam. 

Tylonycteris pachypus aurex Thomas, 1915 

1915. Tylonycteris aurex Thomas, Ann. Mag. N.H. 75.- 228. Astoli, Belgaum, south of 
Bombay, India. Range: Dharwar, Kanara, Coorg, in Peninsular India. 

Tylonycteris robustula Thomas, 1915 

Approximate distribution of species: Yunnan, China; Laos and Annam, Indo- 
China; Malay Peninsula, Sumatra, Java, Borneo, Bali, Celebes, Timor. 

Tylonycteris robustula Thomas, 1915 

1915. Tylonycteris robustula Thomas, Ann. Mag. N.H. 75.- 227. Upper Sarawak, 

Genus BARBASTELLA Gray, 1821 

182 1. Barbastella Gray, London Med. Repos. 75.- 300. Vespertilio barbastellus Schreber. 
1839. Synotus Keyserling & Blasius, Arch. Naturgesch. 5, i : 305. Vespertilio barba- 
stellus Schreber. 

2 species : Barbastella barbastellus, page 1 75 
Barbastella leucomelas, page 176 

Two closely allied species are currently admitted. We follow the classification of 
Tate, 1942, Bull. Amer. Mus. N.H. 80: 264-265, but a change of name is necessary. 
Tate made leucomelas a race of darjelingensis, but the former antedates the latter. 

Barbastella barbastellus Schreber, 1774 Barbastelle 

Approximate distribution of species: England, France, Switzerland, Spain, Italy 
(Ognev), Norway, Sweden, Germany, Holland, Denmark, Poland; Russia (Ukraine, 
Crimea, Caucasus and Transcaucasia). 

Barbastella barbastellus Schreber, 1774 

1774. Vespertilio barbastellus .Schreber, Saugeth. 7.- pi. 55 (text, p. 168). Burgundy, 

1776. Vespertilio barbastelle Muller, Natursyst. Suppl. Regist. Band, 17. Burgundy, 

1836. Barbastellus daubentonii Bell, Hist. Brit. Quad. 7; 63. Burgundy, France. 
1838. Barbastellus communis Gray, Mag. Zool. Bot. 2: 495. Renaming of barbastellus. 



Barbastella leucomelas Cretzschmar, 1826 

Approximate distribution of species: Sinai; Caucasus, Transcaucasia, Russian 
Turkestan (regions of Tashkent and Murgab Oasis), Chinese Turkestan (Yarkand), 
Yunnan, Szcchuan and Kansu, in China; Hondo, Japan; Nepal, Punjab, Sikkim, 
Bhutan Duars, Rajputana (also Gilgit and Assam, according to Blanford) ; Indo- 

Barbastella leucomelas leucomelas Cretzschmar, 1826 

1826. Vespertilio leucomelas Cretzschmar, in RUppcll, Atlas Reise nordl. Afrika, 
Siiugcth. 73,^1. 28b. Arabia Petraea (= Sinai). 

Barbastella leucomelas darjelingensis Hodgson, 1855 

1855. Plecotiis darjelingensis Hodgson, in Horsfield, Ann. Mag. N.H. 16: 103. Darjeel- 
ing, North-Eastern India. (^Vroughton and Tate gave Nepal.) 

1875. Barhastellus dargelinensis Dobson, Proc. Asiat. Soc. Bengal, 85. 

1908. Barbastella barbastella caspica Satunin, Mitt. Kaukas. Mus. 4: 43, 104. Kubaly, 
River Pirsagat, Transcaucasia. 

1916. Barbastella walteri Bianchi, Annu. Mus. Zool. Acad. St. Petersb. 2i: Ixxw 

1916. Barbastella blanfordi Bianchi, loc. cit. Renaming o[ darjelingensis. 

Range: as in the species, except Sinai. 

Genus NYCTICEIUS Rafinesque, 181 9 

1 81 9. JVvcticeius Rafinesque, J. Physique, 88: 417. Nycticeius humeralis Rafinesque 
from North America. 

1824. Mvelicejus Temminck, Mon. Mamm. /.■ xviii. 

1827. Nycticeus Lesson, Man. Mamm. 98. 

1830. Nycticeyx\S^a.g\er, Nat. Syst. Amph. 13. 

1875. Scoteinus Dobson, P.Z.S. 371. Nvcticejus emarginatus Dobson. Valid as a sub- 

3 species in the area covered by this list: 

Nycticeius emarginatus, page 177 
Nycticeius pallidus, page 177 
Nycticeius schlieffeni, page 177 

Hollistcr, 19 1 8, Bull. U.S. Nat. Mus. gg: 93, stated that the Old World species of 
bats, usually placed in Scoteinus, did not seem to differ generically from the American 
species of Nycticeius, and Simpson (1945, 59) places Scoteinus in Nycticeius. We follow 
these authors. .A^. emarginatus is larger than the other two species referred here. A 
comparison of these can be obtained from Dobson (1878), who placed them in 
different genera. But Miller (1907) considered them congeneric. 


Subgenus SCOTEINUS Dobson, 1875 

Nycticeius schliefiTeni Peters, 1859 SchliefFen's Bat 

Approximate distribution of species: Arabia, Egypt, Sudan, Abyssinia, Kenya, 
Eastern Congo, south to South- West Africa and Portuguese East Africa. 

Nycticeius schlieffeni schlieffeni Peters, 1859 

1859. Nycticejus schlieffenii Peters, Mber. Preuss. Akad. Wiss. 224. Cairo, Egypt. 

Nycticeius (?) schlieffeni bedouin Thomas & Wroughton, 1908 
1908. Scoteinus bedouin Thomas & Wroughton, P.Z.S. 540. Lahej, Aden, South- 
western Arabia. 

Nycticeius pallidas Dobson, 1876 Yellow Desert Bat 

Approximate distribution of species: Northern Sind and Punjab, India. 

Nycticeius pallidus Dobson, 1876 

(?) 1834. Vespertilio noctulinus Geoffrey, in Belanger, Voy. aux Indes-Orientales. . . . 

Zool, 92, pi. 3. Bengal. This, if identifiable, may be the first name for 

pallidus. (See Tate, 1942, 282.) 
1876. Scotophilus pallidus Dobson, Monogr. Asiat. Chiroptera, Appendix D, 186. 

Mian Mir, near Lahore, Punjab, North-Western India. 

Nycticeius emarginatus Dobson, 1871 Large-eared Yellow Bat 

Approximate distribution of species: thought to be from some part of India. 

Nycticeius emarginatus Dobson, 1871 

1 87 1. Nycticejus emarginatus Dobson, Proc. Asiat. Soc. Bengal, 211. ? India. 

Genus SCOTOMANES Dobson, 1875 
1875. Scotomanes Dobson, P.Z.S. 371. Nycticejus ornatus Blyth. 
I species: Scotomanes ornatus, page 177 

Scotomanes ornatus Blytli, 1851 Harlequin Bat 

Approximate distribution of species: Szechuan, Yunnan, eastwards to Fukien and 
adjacent states in Southern China; Sikkim, Bengal, Assam, perhaps Northern 

Scoto.manes ornatus ornatus Blyth, 1851 

1851. Nycticejus ornatus Blyth, J. Asiat. Soc. Bengal, 20: 517. Cherrapunji, Khasi 

Hills, Assam. 
1855. Nycticejus nivicolus Hodgson, in Horsfield, Ann. Mag. N.H. 16: 104. Northern 

region of Sikkim Himalayas. 


p.\i.aearc:tic; and Indian mammals 1758-1946 


192 1. Scotomanes ornatus sinensis Thomas, J. Bombay X.H. Soc. :?/■' 772- Kuatun, 
North-\\'estern Fukicn, China. Range: recorded from Szechuan, Hunan, 
Kwangsi, Kwantung, Fokien, Southern China. 


1921. Scotomanes ornatus imhrensis Thomas, J. Bombay N.H. Soc. 2y: 772. Khonsh- 
nong, Jaintia Hills, 3,000 ft., Assam. 

Genus SCOTOPHILUS Leach, 1821 

if')2i. Stntnpliilii\ Leach, 'Lrans. Linn. Soc. London, ij: 69. Seotophiliis kuhlti 

1 83 1. Pachrotus Gray, Zool. Misc. No. i, 38. Seotophiliis kuhlii Leach. 
(?) 1942. Parascotomancs Bourret, C.R. Conseil Rech. Sci. Lidochine, 1^42, 2: 23. 

Scotomanes i Parascotomancs) heaidicui Bourret. 

2 species in the area covered by this list: 
Scotophilus heathi, page 1 79 
Seotophiliis tcmmincki, page 1 78 

The earliest name in this genus is .S'. iu«iita Schreber, 1774, from Senegal. It has a 
vsidc range in Tropical Africa, but we ha\e not heard of its being recorded from 
Piilaearctic Africa. It is, from Dobson's notes, not very widely removed from the 
Indomalayan species. 

Tate, 1942, Bull. Amer. Mus. X.H. 80: 283, reviews the Indomalayan species at 
some length. The earliest name is Scotophilus kulili Leach, 182 1 (Trans. Linn. Soe. 
London, ij: 72, locality unknown). Tate is, hovve\er, unable to identify this form 
specifically, and states that it was based on a juvenile specimen. We here follow the 
classification of Tate, who regards two species as valid: a larger and a smaller, 
occurring side by side in parts of their ranges. We accept Tate's statement that the 
name kuhli is not at the moment certainly identifiable specifically. 

Scotophilus temmincki Horsfield, 1824 Lesser Yellow Bat 

.\ppro.\imate distribution of species: Hainan, Formosa; Ceylon, Peninsula of 
India, where widely distributed, north to Kathiawar, Palanpur, Bengal, Kumaon, 
Sikkim, Bhutan Duars; Mt. Popa, in Burma, Tenasserim; Siam, Annam, in Indo- 
China, Malay States, Java, Bali, Borneo, Philippines. (Bodcnheimer listed "Scoto- 
philus ? temminekr' from Palestine, which is far r>ut of its normal range.) 

SfioTOPHius TEMMINCKI TEMMINCKI Horsficld, 1 824. Extralimital ) 

182 j. Vt\peilUui Iciiniuiicki Horsfield, Zool. Res. Ja\a, fB). Western Ja\a. 


1838. Scotophilus caslamiis Gray, Mag. Zool. Bot. 2: 498. Malacca. Range includes 
Borneo, Annam and Tenasserim. 



1897. Scotophilus wroughtoni Thomas, J. Bombay N.H. Soc. //; 275. Kim, Surat 
district, Western India. Range: Ceylon and India, as above, east to Mt. 
Popa, Burma. 

Scotophilus temmincki consobrinus J. Allen, 1906 

(?) i860. Nycticejus (?) swinhoei Blyth, J. Asiat. Soc. Bengal, 2g: 88. Amoy, Southern 

1906. Scotophilus castaneus consobrinus J. Allen, Bull. Amer. Mus. N.H. 22: 485. 

Rintoi, Island of Hainan. 
Range includes Formosa. 

Scotophilus temmincki gairdneri Kloss, 191 7 

1917. Scotophilus gairdneri K.\oss,}.l>i.Yi. Soc. Siam, 2: 284. Paknampo, Central Siam. 

Scotophilus heathi Horsfield, 1831 Greater Yellow Bat 

Approximate distribution of species (as understood by Tate (1942), i.e. containing 
both the very large and the medium-sized Indomalayan Scotophilus) : Yunnan (and 
possibly parts of South-Eastern China), Hainan; Burma, Bhutan Duars, Sikkim, 
Bengal, Kumaon, Central Provinces, Cutch, Sind, Palanpur, Rajputana, Bombay, 
Peninsular India generally, to Ceylon; Kashmir; Tonkin and Annam, in Indo-China, 
Lower Siam, and evidently Celebes. 

Scotophilus heathi heathi Horsfield, 1831 

1831. Nycticejus heathii Horsfield, P.Z.S. 113. Madras, India. Range includes Raj- 
putana and Ceylon (Tate). 

Scotophilus heathi belangeri I. Geoffroy, 1834 

1B34. Vespertilio belangeri Geoffroy, in Belanger, Voyage aux Indes-Orientales, Zool. 

87. Towns near Pondicherry, Coromandel coast, India. 
1851. Nycticejus luteus Blyth, J. Asiat. Soc. Bengal, 20: 157. "Bengal; Coromandel, 

1851. Scotophilus flaveolus Horsfield, Cat. Mamm. Mus. E. Ind. Co. 37. "Many parts 

of Continental India." 

Probably Scotophilus kuhli of Wroughton's Indian Mammal Survey summary 
should be referred here, but in view of Tate's recent classification of the genus, 
revision of Indian specimens is much needed. 

Wroughton quoted kuhli from Ceylon (but he did not quote heathi from there), 
many localities in Peninsular India, Bengal, Sind, Cutch, Palanpur, Central Pro- 
vinces, Kumaon, Sikkim, Bhutan Duars, \Vestern, Eastern and Central Burma. 
Other localities are Yunnan {kuhli of G. Allen, 1938), Siam, and Tate quoted a 
specimen of fairly similar size from Kashmir. 

Scotophilus heathi insularis J. Allen, 1906 

1906. Scotophilus kuhlii insularis J. Allen, Bull. Amer. Mus. N.H. 22: 485. Rintoi, 
Island of Hainan. 


palaearc:tic; and Indian mammals 1758-1946 

(?) Scotophilus incertae scdis. 

(?) Scotophilus beaulieui Bourret, 1942 

1942. Scotomanes [Parascotomanes) beaulieui Bourret, C.R. Couseil Rech. Sci. Indo- 
chine, IQ^2, 2: 23. Tran-Xinh, Indo-Clhina. 

Genus OTONYCTERIS Peters, 1859 
1859. Otonycteris Peters, Mber. Preuss. Akad. Wiss. 223. Otonycteris hemprichii Peters. 
I species : Otonycteris hemprichi, page 1 80 
There is probably only one valid species in this genus. 

Otonycteris hemprichi Peters, 1859 Hemprich's Long-eared Bat 

Approximate distribution of species: Russian Turlcestan (from Turkmenia to the 

Hissar-Alai, Western Tianshan and Pamir Mountains) ; Persia, Iraq, Palestine 

(Asia Minor, according to Kuznetzov), Arabia; Kashmir (Gilgit); Egypt to Algeria. 

Otonycteris he.mprichi hemprichi Peters, 1859 

1859. Otonycteris hemprichii Peters, Mber. Preuss. Akad. Wiss. 223. No locality. 

(Hemprich & Ehrenberg's collection; probably from some part of North- 

Eastern Africa.) 
1866. Plecotus ustus Fitzingcr & Heuglin, S.B. Akad. Wiss. Wien, 5^, i : 546. Wadi 

Haifa, in Baten-el-Hadjar, Egypt. Norn. nud. 
1873. Plecotus leucophaeus Severtzov, Mem. Soc. Amis. Sci. Nat. Moscou, 8, 2: 18. 

N.W. Turkestan. See also Severtzov, 1876, Ann. Mag. N.H. 18: 42. 
1873. Plecotus auritus brevimanus Severtzov, Mem. Soc. Amis. Sci. Nat. Moscou, 8, 2: 

79. See also Ann. Mag. N.H. 18: 42, 1876. Nee ]enyn%, 1829. 
(?) 1902. O(tonycteris) petersi Anderson & de Winton, Zool. Egypt, Mamm. 120, 

pi. 18, fig. 3. Fao, Persian Gulf. Status ^(/c Ognev. 
1936. Plecotus auritus saharae Laurent, Bull. Soc. Hist. Nat. Afr. N. 2j: 408. El Golea, 

Range: Russian Turkestan, Gilgit, Palestine, Egypt to Algeria. 

Otonycteris (?) hemprichi cinerea Satunin, 1909 

1909. Otonycteris emereus Satunin, Mitt. Kaukas. Mus. ^: 281, 297. Village of 
Nukendzaga, District of Gc, Persian Baluchistan. 

Otonycteris (?) hemprichi jin Cheesman & Hinton, 1924 

1924. Otonycteris jin Cheesman c& Hinton, Ann. Mag. N.H. 14: 549. Hufuf Town, 
Hasa, Arabia. 

Genus PLECOTUS GeofTroy, 18 18 

(?) 1816. Macrotus Leach, Cat. Mamm. & Birds B.NL 5, nam. nud. Macrotus europaeus 

1818. Plecotus Geoffroy, Description de I'Egypte, 2: 1 12. Vespertilio auritus, Linnaeus. 

I species: Plecotus auritus, page 181 



Plecotus auritus Linnaeus, 1758 Long-eared Bat 

Approximate distribution of species: Britain, Ireland, France, Spain, Italy, 
Switzerland, Sweden, Norway, Denmark, Holland, Germany, Yugoslavia, Czecho- 
slovakia, Finland, Poland; Russia, from about 60-62° N., south to the Caucasus, east 
across Siberia to Kamtchatka and Sakhalin, Russian Turkestan; Japan, Kashgar 
(Chinese Turkestan), Tsaidam, Mongolia, China (states of Chihli, Kansu, Szechuan) ; 
Kashmir, Punjab, Kumaon, Nepal; Palestine, Persia, according to Kuzyakin; Egypt 
to Northern Sudan, Tunis, Algeria; Teneriffe (Canary Islands). 

Tate, 1942, Bull. Amer. Mus. N.H. 80: 231, suggests there are three species in 
Eurasia: P. auritus (with races homochrous (synonym puck) and sacrimontis (synonym 
ognevi); P. ariel (with race wardi (synonym kozlovi) ), and P. mordax. 

Plecotus auritus auritus Linnaeus, 1758 

1758. Vespertilio auritus Linnaeus, Syst. Nat. loth ed. /.• 32. Sweden. 

1816. Macrotus europaeus Leach, Cat. Spec. Indig. Mamm. etc. B.M. 5, nom. nud. 

1825. Vespertilio otus Boie, Isis, Jena, 1206. Copenhagen, Denmark. 

1826. Vespertilio cornutus Faber, Isis, Jena, 515. Jutland, Denmark. 

1827. Plecotus communis Lesson, Man. de Mamm. 95. France. 

1829. Plecotus brevimanus Jenyns, Trans. Linn. Soc. London, 16: 55. Grunty Fen, Isle 

of Ely, Cambridgeshire, England. 
1829. Plecotus vulgaris Desmarest, Faune Fran^aise (19) Mamm. 18. France. 
1829. Vespertilio auritus austriacus Fischer, Synops. Mamm. 117. Vienna, Austria. 
(?) 1832. Plecotus peronii I. Geoffroy, Mag. Zool. Paris, 2, i : 2 (not numbered), 

pis. 2-3. 
1832. Plecotus velatus I Geoffroy, Mag. Zool. Paris, 2, i, pi. 2, p. 5 fnot numbered), 

(?) 1838. Plecotus bonapartii Gray, Mag. Zool. Bot. 2: 495, nom. nud. 
1840. Plecotus megalotos Schinz, Europ. Fauna, /.• 19. 
i860. Plecotus kirschbaumii Koch, Ber. Oberhess. Ges. Nat.-u. Heilk. 8: 40. Dillen- 

berg, Oberhessen, Germany. 
1863. Plecotus auritus var. typus Koch, Jb. Nassau. Ver. Naturk. 18: 406. Wiesbaden, 

Nassau, Germany. 
1863. Plecotus auritus var. montanus Koch, loc. cit. \\'esterwald, Nassau, Germany. 
1863. Plecotus auritus var. brevipes Koch, loc. cit. 407. Substitute for kirschbaumii. 
Range: Europe, Siberia, eastwards to Kamtchatka and Sakhalin. 

Plecotus auritus christiei Gray, 1838 

1829. Vespertilio auritus aegjptius Fischer, Synops. Mamm. 117, not Vespertilio pipi- 

strellus var. aegjptius Fischer, ibid. 105. 
1838. Plecotus christii Gray, Mag. Zool. Bot. 2: 495. North Africa. 
1878. Plecotus aegyptiacus "I. Geoff.", Dobson, Cat. Chiroptera B.M. 178. Egypt. 

(See Thomas, 1911, P.Z.S. 160.) 
Range: Egypt, to Aswan and Northern Sudan; Tunis; Palestine. 

Plecotus auritus homochrous Hodgson, 1847 

1847. Plecotus homochrous Hodgson, J. Asiat. Soc. Bengal, 16: 895. Nepal. Ranges to 

PALAKARCIR: and IXUIAX mammals 1758-1946 

Plecotus auritus teneriffae Barrett-Hamilton, 1907 

1907. Plecotus temriffac Barrett-Hamilton, Ann. Mag. N.H. 20: 520. Orotava, Island 
of Tcncrifle. 

Plecotus .vuritus puck Barrett-Hamilton, 1907 

1907. Plecotus puck Barrett-Hamilton, Ann. Mag. N.H. 20: 521. Murrce, 7,500 ft., 

Punjab, India. ? Synonym of homocfnous (Tate). 

Plecotus auritus sacr:montis G. Allen, 1908 

1908. Plecotus sacrimontis G. Allen, Bull. Mus. Comp. Zool. Harvard, 52.- 50. Mt. 

Fuji, Japan. 
1927. Plecotus auritus ognevi Kishida, Zool. Mag. Tokyo, 59.- 418. North Sakhalin. 

Plecotus auritus wardi Thomas, 191 1 

191 1. Plecotus wardi Thomas, Ann. Mag. N.H. 7; 209. Leh, Ladak, Kashmir. 

Range includes high parts of the Gaucasus, according to Kuzyakin, also 

Russian Turkestan; and probably Zungaria. 

Plecotus auritus ariel Thomas, 191 1 

1911. Plecotus ariel Thomas, Abstr. P.Z.S. 3; P.Z.S. 160. Tatsienlu, 8,400 ft., 
Szechuan, Ghina. 

Plecotus auritus kozlovi Bobrinskii, 1926 

1926. Plecotus auritus kozlovi Bobrinskii, C.R. Acad. Sci. U.R.S.S., A, 98. Barun 

Zasak, Eastern Tsaidam, Ghincsc Gentral Asia. Range includes Gobi, 


Plecotus auritus mordax Thomas, 192G 

1926. Plecotui worrfa.v Thomas, Ann. Mag. N.H. 18: 306. Kashgar, Chinese Turkestan. 

Plecotus auritus mrridionalis Martino, 1940 

1940. Plecotus auritus mendwnaUs Martino, Ann. Mag. N.H. 5: 494. Sueti Miklavz pri 
Ormozu, Slovenia, Yugoslavia. 

Subfamily M i n i o p t e r i n a e 

Genus MINIOPTERUS Bonaparte, 1837 

1837. Minio/iterus Bonaparte, Fauna Ital. /.• fasc. 20, under Vesperlilio cmarginatus. 

I'espertilio ursiuii Bonaparte = Vespertilio ichreihersii Kuhl. 
i860. Miniopteris Gray, Ann. Mag. N.H. ly: 91. 
1892. Mmyoptcrus Winge, jordfundne og nulevcnde Flagermus (Chiroptera) fra 

Lagoa Santa, Minas Geraes, Brasilien, 36. 
1900. Minneopterus Lampe.Jb. Nassau. Ver. Naturk, jj, Catal. Siiugcth. Samml. 12. 

2 species in the area covered by this list: 
Miniopterus australis, page 184 
Miniopterw, ichreihersi, page 183 

In this genus we follow Tate, 1941, Bull. Atner. .\lu\. .\'.//. y8: 568. 


Miniopterus schreibersi Kuhl, 1819 Schreibers' Bat. Long-winged Bat 

Approximate distribution of species: Spain, France, Switzerland, Italy, Germany, 
Hungary, Poland, Sardinia, Montenegro, Bulgaria, Greece, Crete; Crimea, Caucasus 
and Kopet-Dag Mountains, South-West Russian Turkestan; Northern Persia, Pales- 
tine; Japan, Liukiu Islands, Formosa, China (states of Chihli, Chekiang, Fukien, 
etc.), Hainan; Ceylon, Peninsular India (Western Ghats), Kumaon, Nepal, Mt. 
Popa, in Burma; Java, Sumatra, Borneo, Philippine Islands, to New Guinea and 
Northern Australia; Algeria. 

Miniopterus schreibersi schreibersi Kuhl, 181 g 

1819. Vespertilio schreibersii Kuhl, Ann. Wetterau. Ges. Naturk. 4, 2: 185. Kulmbazer 

Cave, mountains of Southern Bannat, Hungary. 
1837. Vespertilio ursinii Bonaparte, Faun. Ital. /.• fasc. 21. Monte Corno, Ascoli, 

1840. Vespertilio orsinii Temminck, Mon. Mamm. 2: 179. Modification ot^ ursinii. 
1926. Miniopterus schreibersii italicus Dal Piaz, Atti Soc. Ven. -Trent. Sci. Nat. 16: 61. 

Arma del Frate, Foligno, near Finalese, Liguria, Italy. 
1936. Miniopterus schreibersii inexspectatus Heinrich, Mitt. Naturw. Inst. Sofia, g: 34. 

Strandja-Balkan, Bulgaria. 

Range: Europe, Algeria. 

Miniopterus schreibersi fuliginosus Hodgson, 1835 

1835. Vespertilio fuliginosa Hodgson, J. Asiat. Soc. Bengal, 4: 700. Nepal. 

1906. Miniopterus schreibersi japoniae Thomas, P.Z.S. igo^, 2: 338. Tano, Miyasaki 

Ken, Kiushiu, 500 ft., Japan. 

1923. Miniopterus schreibersii parvipes G. Allen, Amer. Mus. Nov. No. 85: 7. Yenping, 

Fukien, Southern China. 

Range: Nepal, Ceylon, Southern India, Burma; Fukien and Hunan, in China; 
Hainan; Japan. 

Miniopterus schreibersi blepotis Temminck, 1840 

1840. Vespertilio blepotis Temminck, Mon. Mamm. 2: 212. Java. 

1902. Miniopterus fuscus Bonhote, Nov. Zool. g: 626. Okinawa, Liukiu Islands. 

1924. Miniopterus fuscus rajejamae Kuroda, New Mamm. Riukiu Islands, 6. Ishigaki- 

Mura, Ishigaki, Liukiu Islands. 

Range: Liukiu Islands, also Sumatra, Java, Borneo, Philippine Islands. 

Miniopterus schreibersi pallidus Thomas, 1907 

1907. Miniopterus schreibersii pallidus Thomas, Ann. Mag. N.H. 20: 197. Southern 

shore of Caspian Sea, Northern Persia. Range: to Transcaspia (Ognev). 

Miniopterus schreibersi chinensis Thomas, 1908 

1908. Miniopterus schreibersi chinensis Thomas, P.Z.S. 638. Thirty miles west of Pekin, 

Chihli, North-Eastern China. Range includes Chekiang, China. 



Miniopterus australis Tomes, 1858 

Approximate distribution of species: Nicobar Islands, Madras, India, Hainan. 
Also Java, Borneo, Philippine Islands, Amboina, Loyalty Islands, etc. 

(Miniopterus australis australis Tomes, 1858. Extralimital) 
1858. Miniopterus australts Tomes, P.Z.S. 125. Loyalty Islands (21° S., 167.30' E., 
South Pacific). Ranges to New Guinea. 

Miniopterus aiistralis pusillus Dobson, 1876 

1876. Miniopterus pusillus Dobson, Monogr. Asiat. Chiroptcra, 162. Madras, India 

(Tate). But \Vroughton gave Nicobar Islands as type locality. Range; 

Madras, Nicobar Islands, Hainan and Borneo (Tate). 

Subfamily M u r i n i n a e 
For review, see Tate, 1941, Bull. Arner. Mus. N.H. y8: 575. 

Genus MURINA Gray, 1842 

1842. Murina Gray, .^nn. Mag. N.H. 10: 258. Vespertilio suillus Temminck, from Java. 
1842. Ocvpetes Lesson, Nouv. Tabl. Regne Anim. 30 (part). Not of Wagler, 1832. 
1915. Harpiola Thomas, Ann. Mag. N.H. 16: 309. Murina grisea Peters. Valid as a 

Tate ( 1941, 577) gave a key to the species groups. These, in the region now under 
discussion, amount to five, one of which is subgenerically (or generically) separated 
as Harpiola on account of some dental characters. Far too many species are standing 
in the genus. Tate has shown clearly how the groups can be divided, and until the 
contrary is proved we propose to assume that the other named forms are races 
respectively of the five names listed below: 

Murina aurata, page 184 
Murina eyclotis, page 186 
Murina grisea, page 1 86 
Murina huttoni, page 186 
Murina leucogaster, page 185 

Subgenus MURINA Gray,- 1842 

Murina aurata Milne-Edwards, 1872 Little Tube-nosed Bat 

-Approximate distribution of species: Ussuri district of South-Eastern Siberia, 
Japan, Szcchuan and Yunnan (in China), Sikkim, Burma. 

MuRiXA AURATA AURATA Milne-Edwards, 1872 

1872. Murina aurata Milne-Edwards, Rech. H.N. Mamm. 250, pi. 37b, fig. i ; pi. 37c, 

fig. 2. Moupin, Szcchuan, China. Ranges to Yunnan, China, and Sikkim 

according to Wroughton. 
1907. Murina aurila .Miller, Bull. U.S. Nat. Mus. f^j: 230. 



1891. Harpiocephalus feae Thomas, Ann. Mus. Stor. Nat. Genova, lo: 884; 926-927 
(1892). Biapo, Karen Hills, Burma. 


1913. Murina ussuriensis Ognev, Annu. Mus. Zool. Acad. St. Petersb. i8: 402. 
Evseevka, Imansky district, Ussuri and Odarka, Chanka Lake, Ussuri 
district, South-Eastern Siberia. Widely distributed in Japan. 

Murina leucogaster Milne-Edwards, 1872 Great Tube-nosed Bat 

Approximate distribution of species: Siberia, known from Upper Yenesei, Kuz- 
netzk Ala-Tau, Lake Teletzkoie in Altai, Ussuri region, Sakhalin; Japan, China 
(states of Szechuan and Fukien) ; Manchuria; near Darjeeling, North-Eastern India. 

Murina leucogaster leucogaster Milne-Edwards, 1872 

1872. Murina leucogaster Milne-Edwards, Rech. H. N. Mamm. 252, pi. 37b, fig. i 

{2 in error); pi. 37c, fig. 3. Moupin district, Szechuan, China. 
1899. Murina leucogastra Thomas, P.Z.S. i8g8: "jii. 
Ranges to Fukien, China. 

Murina leucogaster hilgendorfi Peters, 1880 

1880. Harpyocephalus hilgendorfi Peters, Mber. Preuss. Akad. Wiss. 24. Near Tokyo, 
Yeddo (= Hondo), Japan. 

Murin.a. leucog,\ster sibirica Kastschenko, 1905 

1905. Harpiocephalus leucogaster sibiricus Kastschenko, Observ. Mamm. \V. Siberia & 
Turkestan, 102b. Tomsk region, Siberia. (Kuzyakin, in Bobrinskii (1944), 
ignores this name and uses hilgendorfi for the Siberian representative of this 

Murina leucogaster ognevi Bianchi, 19 16 

1916. Murina ognevi Bianchi, Annu. Mus. Zool. Acad. St. Petersb. 21: ixxviii. 
Vladivostock, Eastern Siberia. Remarks as for last race. 

Murina leucogaster rubex Thomas, 1916 

1916. Murina rubex Thomas, J. Bombay N.H. Soc. 24: 639. Pashok, near Darjeeling, 
North-Eastern India. 

Murina leucogaster fusca Sowerby, 1922 

1922. Murina huttonii fuscus Sowerby, J. Mamm. 3: 46. Northern Kirin, Manchuria. 

Murina leucogaster intermedia Mori, 1933 

1933. Murina hilgendorfi intermedia Mori, J. Chosen N.H. Soc. 16: 2, 5. Mt. Kongo, 

P.\I.AF,.\RC:TIC: and IXDIAX mammals i 758-1946 

Murina huttoni Peters, 1872 

Approximate distribution of species: Kashmir, Kumaon, Sikkim district. Western 
Burma, Fukicn (China), Tonkin and Laos (Indo-China). 


1872. Harpyiocephalus huttoni Peters, Mber. Preuss. Akad. VViss. 257. P.Z.S. 711. 
Dehra Dun, Kumaon, North-Wcstern India. Also recorded Irom DarjeeHng. 

Murina (?) huttoni tubinarjs Scully, 1881 

1 88 1. Harpiocephalus lubinaris Scully, P.Z.S. 200. Gilgit, Kashmir. Has also been 

recorded from Tonkin and Laos by Osgood, and from DarjeeHng and Clhin 

Hills. Tate states (1941, 577) "^huttoni ( -^ tuhimuis?)" . 

Murina huttoni rubella Thomas, 1914 

1914. Murina huttoni rubella Thomas, Ann. NLag. N.H. 13: 440. Kuatun, Fukien, 
South-Eastcrn China. 

Murina cyclotis Dobson, 1872 

Approximate distribution of species: Hainan; Sikkim, Western and Northern 
Burma; Tonkin and Laos, Indo-China; Clcylon. Recorded also from the Philippine 

Murina cyclotis cyclotis Dobson, 1872 

1872. Murina cyclotis Dobson, Proc. Asiat. Soc. Bengal, 210. Darjecling, xNorth- 
Eastern India (Tate). Range includes Burma, Indo-China, Hainan. 

Murina cyclotis eileenae Phillips, 1932 

1932. Murina eileenae Phillips, Ceylon J. Sci., B, iG, 3: 329. Mousakande, Gamma- 
duwa, 3,000 ft., Ceylon. 

Ineertae sedis 

Murina puta Kishida, 1924, Zool. Mag. Tokyo, 36: 30-49, 127-139. Formosa. 


Subgenus HARPIOLA Thomas, 191 5 

Murina grisea Peters, 1872 Peters' Tuhc-nosed Bat 

Approximate distribution of species: Kumaon, North- Western India. 

MuRiN.^i GRISEA Peters, 1872 

1872. Mimna i^risea Peters, Mber. Preuss. Akad. Wiss. 258. P.Z.S. 712. Jeripanee, 
Mussoori, 5,50ij ft., Kumaon, North-Wcstern Himalayas. 



Genus HARPIOCEPHALUS Gray, 1842 

1842. Harpiocephalus Gray, Ann. Mag. N.H. 10: 259. Harpiocephalus rufus Gray = 

Vespertilio harpia Temminck. 
1866. Harpyiocephalus Gray, Ann. Mag. N.H. ly: 90. 

For characters of this genus compared with Murina, see Miller, 1907, Families and 
Genera of Bats, 229. 

I species: Harpiocephalus harpia, page 187 

Harpiocephalus harpia Temminck, 1840 Hairy- winged Bat 

Approximate distribution of species: Formosa; Darjeeling, Bhutan Duars, Palni 
Hills, in Southern India, Northern Burma; Indo-China; Sumatra, Java; Amboina 

Harpiocephalus harpia harpia Temminck, 1840 

1840. Vespertilio harpia Temminck, Mon. Mamm. 2: 219, pi. 55. Mt. Gede, Java. 
1842. Harpiocephalus rufus Gray, Ann. Mag. N.H. 10: 259. New name for harpia. 
(?) 1858. Vespertilio pearsonii Tomes, P.Z.S. 87. Locality unknown. 
Recorded from Formosa (Kuroda). 

Harpiocephalus harpia l.^syurus Hodgson, 1847 

1847. Noctulinia lasyura Hodgson, J. Asiat. Soc. Bengal, 16: 896. "Central Hills, sub- 
Himalayas." Darjeeling, according to Wroughton. Also occurs in Bhutan 

Harpiocephalus harpia rufulus G. Allen, 1913 

1913. Harpiocephalus rufulus G. Allen, Proc. Biol. Soc. Wa.shington, 26: 214. Lao-Kai, 
Tonkin, Indo-Chiha. 

Harpiocephalus harpia madrassius Thomas, 1923 

1923. Harpiocephalus harpia madrassius Thomas, J. Bombay N.H. Soc. 2g: 88. Perumal, 
Palni Hills, Southern India. 

Harpiocephalus (?) harpia mordax Thomas, 1923 

1923. Harpiocephalus mordax Thomas, J. Bombay N.H. Soc. 2g: 88. Mogok, Upper 

Subfamily Kerivoulinae 

Genus KERFVOULA Gray, 1842 

1842. Kerivoula Gray, Ann. Mag. N.H. 10: 258. Vespertilio pictus Pallas (Peters, 1866). 
1849. Kirivoula Gervais, Diet. Univ. H.N. ij: 213. 
1 891. Cerivoula Blanford, Fauna Brit. Ind. Mamm. 338. 

1905. Phoniscus Miller, Proc. Biol. Soc. Washington, 18: 229. Phoniscus alrox Miller. 
Valid as a subgenus. For status see Simpson ( 1 945, 60) and Tate ( 1 94 1 , 586) . 

N . 187 


The Oriental members of the genus were reviewed by Tate (1941, 584). Dobson 
(1878, 331) also gave a key to the African and Asiatic species then known. 

3 species in the area covered by this list : 
Kcrivoula hardwickei, page 188 
Kerh'oula papulosa, page 1 89 
Kerivoula picta, page 1 88 

Kerivoula picta Pallas, 1 767 Painted Bat 

Approximate distribution of species; Kwantung, in Southern China, Hainan; 
Ceylon and Southern -India (Western Ghats, Dharwar). (Blanford gave several other 
Indian localities, including Sikkim, Bengal and Burma.) Malay States, Sumatra, 
Java, Bali, Borneo. 

Kerivoula picta picta Pallas, 1767 

1767. Vespeiiilio piclus Pallas, Spic. Zool. j: 7. Most authors cite Peninsular India as 
the type locality. Tate, however, thinks it came from Ternate, Moluccas 
(near Halmahcra). 
1832. Vespertilio kirivoida Cuvier, Nouv. Arch. Mus. H.N. Paris, /.• 9. 
Range includes Malay States, eastwards to Bali and Borneo; also Ceylon and 
Southern India. 

Kerivoul.\ pict.\ belllssima Thomas, 1906 

1906. Ktrivoula picta bellissima Thomas, Ann. Mag. N.H. //.• 423. Pakhoi, Southern 
Kwantung, Southern China. Range includes Haindn. 

Kerivoula hardwickei Horsfield, 1824 Hardwicke's Bat 

Approximate distribution of species: Szechuan, Kwangsi and Fukien in Southern 
China; Darjeeling, Mysore in Southern India, Ceylon, Burma, (Blanford also 
quoted Assam and the Punjab); ? Indo China; Malay States, Mentawei Islands 
west of Sumatra, Ja\a, Bali, Borneo, Celebes and probably represented in the 
Philippine Islands. 

Kerivoula hardwickei hardwickei Horsfield, 1824 
1824. Vespeiiilio liaidwickii Horsfield, Zool. Res. Ja\a, (K). J.i\-.i. 
1 87 1. Kerivoula fusca Dobson, Proc. Asiat. Soc. Bengal, 215. No locality. 
Range: apparently includes Darjeeling, as well as Malay States, Borneo, Java, Bali, 

Kerivoula hardwickei depressa Miller, 1906 

1906. Kerivoula depressa Miller, Proc. Biol. Soc. Washington, ig: 64. Biapo, Karin 
Hills, Southern Burma. Range includes Szechuan and Fukien, China. 

Kerivoula hardwickei crypta Wroughton & Ryley, 19 13 

1913. Kcrivoula crypta Wroughton & Ryley, J. Bombay N.H. Soc. 22: 14. Shimoga, 

Mysore, Southern India. Range includes Upper Burma (Kaulback Coll., 



Kerivoula hardwickei malpasi Phillips, 1932 

1932. Kerivoula malpasi Phillips, Ceylon J. Sci., B, 16: 331. Kumbalgamuwa, 3,000 ft., 
Mulhalkelle district, Central Province, Ceylon. 

Kerivoula papillosa Temminck, 1840 

Approximate distribution of species: Calcutta, India; Indo-China; Malay States, 
Sumatra, Java, Borneo. 

(Kerivoula papillosa papillos.a Temminck, 1840. Extralimital) 

1840. Vespertilio papillosa Terriminck, Mon. Mamm. 2: 220. Bantam, Java. 

Kerivoula papillosa lenis Thomas, 19 16 

1916. Kerivoula lenis Thomas, J. Bombay N.H. Soc. 24: 416. Calcutta, Bengal, India. 

Kerivoula papillosa malayana Chasen, 1940 

1940. Kerivoula papillosa malayana Chasen, Bull. Raffles Mus. 75.- 55. Gintang Bidei, 

Selangor-Pahang boundary, 2,300 ft., Malay States. Recorded from Tonkin, 

Indo-China (Tate, 1947). 


(Not including Family Hominidae) 

FAMILIES: Cercopithecidae, page 192 
Lorisidae, page 190 
Pongidae, page 2 1 1 

Simpson, 1945, also refers the family Tupaiidae to the Primates, and discusses this 
classification at length (pp. 176, 182, 183). It is by no means conclusively proved, 
however, that this classification is the correct one, and for the present we prefer to 
regard them as belonging to the order Insectivora. It appears to us that of the 
Primates the more specialized members, the Anthropoidea, are easily defined and 
distinguished from the lower orders of Mammalia, such as the Insectivora, but that 
the more generalized members, the Prosimii of Simpson (perhaps excepting the 
Tarsiidae) are not so easily separable from the lower orders. We would particularly 
draw attention to Simpson's amusing explanation (pp. 180, 181) of the confusion 
which exists in this order, particularly as regards nomenclature. 

Special works of reference include Elliot, 1913, a Review of the Primates, Monogr. 
Amer. Mus. M.H., 3 volumes, in which there is wholesale splitting, but which remains 
the best single source of information on living Primates; and Pocock, 1939, Fauna 
British India, Mammalia, i: 13, which gives a classification of the Indian Primates 
and clears up a great deal of former nomenclatural difficulty. The Malaysian forms 
are listed, in apparently good order, by Chasen (1940). Pocock, 1934, P.^-S. 895, 
reviewed the Langurs, and 1927, P-Z-S- 719, the Gibbons. He also published several 
short papers on Macaques. 



Apart from the Hominidae and, as explained above, the Tupaiidae, Simpson 
(1945) classified the Indian and Palaearctic recent Primates as follows: 

Suborder: PROSIMII 

Infraorder: Lorisiformes 
Family: Lorisidae 


Superfamily : Cercopithecoidea 

Family: Ccrcopithccidae (with subfamilies Cercopithecinae and 

Superfamily : Hominoidea (in part) 

Family: Pongidae (subfamilies Hylbbatinae and Ponginae (extra- 
limital) ) 



Genera: Loris, page 190 

Js'yclicebus, page 19I 

For generic characters, see Pocock, 1939, Fauna British India, Mamm. i: 165. 

Genus LORIS E. Gcoftroy, 1796 

1785. Tardigradus Boddaert, Elench. Anim. 43, 67. Tardigradus Inris Boddaert = 
Lemur tardigradus Linnaeus. Not of Brisson, 17G2. 

1796. Loris E. Geoffroy, Mag. Encycl. /.• 48. Loris gracilis Geoffroy = Ltnmr tardi- 
gradus Linnaeus. 

181 1. Slenops lUiger, Prodr. Syst. Mamm. ct. Avium, 73. Lemur tardigradus Linnaeus. 

1 species: Loris tardigradus, page 190 

Loris tardigradus Linnaeus, 1 758 Slender Loris 

Approximate distribution of species: Ceylon and Southern India (Eastern Ghats, 
Mysore, Malabar, Travancore, Coorg). 

Loris tardigradus tardk;radus Linnaeus, 1758 

i7-,8. Lemur tardigradus Linnaeus, Syst. Nat. loth ed. /.■ 29. Ceylon. 

1706. Loris gracilis E. Geoffroy, Mag. Encycl. /.■ 48. Ceylon. 

1804. Loris ceylonicus Fischer, Anat. Maki, /.• 28. Ceylon. 

1004. Lurts gracilis zeylanieus Lydekker, P.Z.S. 2: 346. Peradcniya, Ceylon. (See 

Pocock, 1939, 181.) 
Range: low-country wet zone ot Ceylon. 



1908. Loris lydekkerianus Cabrera, Bol. Soc. Esp. H.N. Madrid, 139. Madras, India. 
Range: Eastern Ghats, westwards to Mangalore and Mysore, India. 

Loris tardigradus malabaricus \Vroughton, 19 17 

1917. Loris malabaricus Wroughton, J. Bombay N.H. Soc. 23: 45. Huvinakadu 

Estate, Kutta, South Coorg, 2,843 ft-. India. Range: Malabar district, 

Wynaad, South Coorg, Travancore. 

Loris tardigradus grandis Hill & Phillips, 1932 

1932. Loris tardigradus grandis Hill & Phillips, Ceyl.J. Sci. (B), ij: iii. Mousekanda, 

Gammaduwa, 2,200 ft., Central Province, Ceylon. Range: "Probably 
throughout the lower foothills of the mountain cluster of the Central and 
Uva Provinces," up to 3,500 ft. approximately. 

Loris tardigradus nordicus Hill, 1933 

1933. Loris tardigradus nordicus Hill, Ceyl.J. Sci. (B), iS: 113, 120. Talawa, 50 ft.. 

North Central Province, Ceylon. Range: the dry zone of the North Pro- 
vince, North Central Province and Central Province of Ceylon, from just 
above sea level up to 650 ft. 

Loris tardigradus nycticeboides Hill, 1942 

1942. Loris tardigradus nycticeboides Hill, J. Bombay N.H. Soc. 43: 73. Horton Plains 
6,000 ft., Ceylon. 

Genus NYCTICEBUS E. Geoffroy, 1812 

1812. Nycticebus E. Geoffroy, Ann. Mus. H.N. Paris, ig: 163. Nycticebus bengalensis 

2 species: Nycticebus coucang, page 191 
Nycticebus pygmaeus, page 1 92 

Pocock (1939) thought that there was only one species in this genus, but Osgood 
(1932) lists two forms from Indo-China, and as there is an apparent geographical 
overlap between them and they occur together, pygmaeus is here regarded as a valid, 
smaller species. 

Nycticebus coucang Boddaert, 1785 Slow Loris 

Approximate distribution of species: Assam, Chittagong, Burma, Tenasserim, 
Siam, Indo-China, Malay States, Sumatra, Java, Borneo, and some adjacent small 
islands to Philippine Islands. Possibly into Yunnan. 

Nycticebus coucang coucang Boddaert, 1785 

1785. Tardigradus coucang Boddaert, Elench. Anim. 67. Locality unknown (probably 

Malacca (Chasen) ). Range: Mergui Archipelago (King Island quoted by 

Pocock), Malay States, Sumatra. 



NvcTic.F.Brs i:oii:AN(; bexcalensis Laccpcde, 180(1 

1801). Loii henjJiileNiis Laccpcde, Seances des ficoles normalcs, Tome 8: 68. Bengal. 

1804. Loris bengalensis Fischer, Anat. Maki, /.■ 30. Bengal. 

1867. Nycticebus cinereus Milne-Edwards, Nouv. Arch. Mus. Bull. 3: 9. Bangkok, 
Siam. Although G. Allen and Osgood listed this as a valid race, Pocock says 
it cannot be distinguished from the earlier-named bengalensis. 

(?) 1904. Nycticebiis tardigradus Ijpieus Lydekker, P.Z.S. 2: 345. 

1 92 1. Nrctkebus uicanus Thomas, Ann. Mag. N.H. 8: 627. Kyeikpadeih, Pegu, 

Range: Assam, Ohittagong, Burma (? into Yunnan), Indo-China, Siam. 

Nycticebus coucang tenasserimensis Elliot, 1913 

1913. Nycticebus tenasserimensis Elliot, Rev. Primates, /.• 25. Amherst, Northern 
Tenasserim. Range: Tenasserim and South-Western Siam. 

Nycticebus pygmaeus Bonhote, 1907 Lesser Slow Loris 

Approximate distribution of species: Indo-China. 

Nycticebus pygmaeus Bonhote, 1907 

1907. Nycticebus pvgmacus Bonhote, Abstr. P.Z.S. No. 38, 2. P.Z.S. 4. Nhatrang, 
Annam, Indo-China. Osgood (1932) cjuoted this form from Annam, Laos, 
Cochin-China and Tonkin, apparently occurring with N. coucang bengalensis 
{''cinereus") which was quoted from Laos and Annam. 

Suborder Anthropoidea 


Genera: Macaca, page 193 
Papio, page 200 
Presbylis, page 203 
Prgathnx, page 202 
Rhinojnihecus, page 201 

This family is divided into two subfamilies: the Colobinae, which contains the 
Langurs and Leaf-eating Monkeys, Presbytis, Pygathrix, Rhinopithecus; and the Cerco- 
pithecinae, to which Papio and Macaca belong. Some authors gi\e the two divisions 
family rank. 

It is interesting to note that Winge, 1924, Pattedyr Slaegter, 2: 277, recognized only 
five genera in the whole family, which he divided in a different way from that 
usually agreed on: namely, he contrasted a groUp Cercopithecini, with weaker 
cheekteeth, shorter face, containing the African Cercopithecus plus the Langurs and 
Leaf-eating Monkeys Semnopithecus (= the Asiatic genera currently recognized) and 
the African Colobus with a group "Cynocephali" with cheekteeth stronger, face 
longer, containing Macaca and "Cynocephalus" = Papio. 


Subfamily Cercopithecinae 

Genus MACACA Lacepede, 1 799 

1758. Simia Linnaeus, Syst. Nat. loth ed. /; 25. Simla sylvanus Linnaeus. 

(By suspension of the Rules the name Simia is suppressed, see Opinion 1 14 
of Internat. Comm. on Zool. Nomenclature.) 
1799. Alacaca Lacepede, Tabl. Mamm. 4. Simia inuus Linnaeus = Simia sylvanus 

1 81 2. Inuus E. GeofTroy, Ann. Mus. H.N. Paris, ig: 100. Inuus ccaudatus Geoffroy = 

Simia sylvanus Linnaeus. 
1816. Sylvanus Oken, Lehrb. Naturgesch. ^, 2: 1223. Inuus ecaudatus Geoffroy = 

Simia sylvanus Linnaeus. 
1820. Silenus Goldfuss, Handbuch Zool. 2: 479. Cynocephalus silenus Schreber = Simia 

silenus Linnaeus. 
1824. Magotus Ritgen, Nat. Eintheilung Saugeth. 33. "Les Magots" of Cuvier. 

1827. Magus Lesson, Man. Mamm. 43. Magus sylvanus and M. maurus. 

1828. Pithes Burnett, Quart. J. Sci. Lit. & Art. 26, 2: 307. Pithes sylvanus = Simia 

sylvanus Linnaeus. 

1839. Alaimon Wagner, Schreb. Saugeth. Suppl. /.■ iv bis and 141. Inuus silenus = 

Simia silenus Linnaeus. 

1840. Rhesus Lesson, Rev. Zool. 2: 70, nom. nud. 1840, Spec. Mamm. 95. Cercopithecus 

mulatta Zimmermann. 

1 84 1. Salmacis Gloger, Gemeinn. Naturges. /.• 35. New name for Macaca. 

1848. Lyssodes Gistel, Naturgesch. Thier. f. hohere Schulen, 9. Macaca speciosus 
F. Cuvier. 

1862. Vetulus Reichenbach, Vollstand. Nat. Affen, 125. New name for Silenus Lesson. 

1862. Cynamolgus Reichenbach, Vollstand. Nat. Affen, 130. Macacus irus Cuvier {Jide 

1862. ^ati Reichenbach, Vollstand. Nat. Affen, 130. Macaca radiata Geoffroy {fide 

1862. Nemestrinus Reichenbach, Vollstand. Nat. Affen, 139. Macaca nemestrina Lin- 
naeus. Not of Latreille, 1802. 

1913. Pithecus Elliot, Rev. Primates, 2: 176. Not of Cuvier & Geoffroy, 1795. 

Macacus of many earlier authors, including Blanford, 1888, Fauna Brit. India, 
/.• II. 

1 1 species in the area covered by this list : 
Macaca assamensis, page 198 
Macaca cyclopis, page 1 98 
Macaca fuscata, page 199 
Macaca irus, page 196 
Macaca mulatta, page 197 
Macaca nemestrina, page 195 
Macaca radiata, page 195 
Macaca silenus, page 195 
Macaca sinica, page 194 
Macaca speciosa, page 199 
Macaca sylvana, page 200 



The type is the Xorth-W'est African species .\/. sylvana. Various subgcucric names 
are available for some of the other species; Pocock, 1939, Fauna British India, Mam- 
malia, I, gives a key to eight of the above species which occur in India, and lists the 
subgeneric groups. As far as distribution is concerned, three of the species, M. sinica, 
'M. radiata, M. silenus, are confined to Peninsular India and/or to Gcylon; two, M. 
ncnustrina and M. irus, occur together from Burma south-eastwards through the 
Malaysian region covered by Chasen (1940); the species M. mulatto, M. speciosa and 
,\/. assamensis arc roughly Himalayan — Indo-China — Chinese in range; and the 
other two species, Al. fuscata and M. cvclopis, arc from Japan and Formosa respec- 
tively. The genotype, a tailless species, lacks the "cap" of hairs on the head which is 
usually present in the species inhabiting India, mulatta and irus excepted. Pocock 
ip. 33) states that the "cap" is also absent \n fuscata, which is a species with a short, 
hairy tail and appears to be nearly allied to speciosa (although Pocock definitely states 
(p. 70) that speciosa differs from fuscata in the structure of the glans penis); and in 
cvclopis, which probably belongs to the mulatta group, as it seems very like M. 
assamensis. But its tail is about 68 per cent, of the head and body length, according to 
measurements given by Elliot, which is longer than is normal in assamensis, and the 
tail is black and very well haired, which character seems to distinguish from assa- 
mensis in the material examined. 

Macaca sinica group 

The name ^a// Reichenbach, 1862, is available for these species if subgeneric 
division is required. Long-tailed species, diflfering from their allies, according to 
Pocock, in the structure of the male genitalia. 

Macaca sinica Linnaeus, 1771 Toque Monkey 

Approximate distribution of species: Ceylon. 

Mac.-\ca sinica sinica Linnaeus, 1771 

1 77 1. Simla sinica Linnaeus, Mant. Plant. 521. Locality unknown. 

1862. Cynaniolgus ''^ati) audehrrti Reichenbach, X'ollstand. Xat. Affen, 132. 

1863. Macaca pileatus Blyth, Cat. Mamm. Mus. Asiat. Soc. 9. Not of Kerr, 1792. 

1 93 1. Macaca sinica inaurea Pocock, J. Bombay N.H. Soc. 55.' 286. Chcddikulam, 

North Province, Ceylon. 
Range : low-country' dry zone, from extreme north to extreme south of Ceylon. 

Macaca sinica aurifrons Pocock, 1931 

1 93 1. Macaca sinica aurifrons Pocock, J. Bombay N.H. Soc. jjj.- 286. Rayigam Korale, 

Western Province, Ceylon. Range: low-country wet zone and central hill 

zone of Ceylon. 

M.-\cac.-\ sink:a opisthomelas Hill, 1942 

1942. Macaca '-Cfl/0 sinica opisthomelas Hill, J. Bombay N.H. Soc. ^j.- 402. Horton 
Plains, Highlands of Ceylon. 



Macaca radiata Geoffroy, 1812 Bonnet Monkey 

Approximate distribution of species: Peninsular India, north to Satara and the 
Godaveri River. Closely allied to and perhaps representing sinica on the mainland. 
For characters see Pocock (1939, 33, 38). 

Macaca r.vdiata radiata Geoffroy, 181 2 

1812. Cercocebus radiatus E. Geoffroy, Ann. Mus. H.N. Paris, ig: 98. Locality un- 
known. Range: Satara, Kanara, Mysore, Coorg, Nilgiri and Palni Hills, 
Cochin, Eastern Ghats, etc., in Peninsular India. 

Macaco radi.\t.'^ diluta Pocock, 193 1 

1931. Macaca radiata diluta Pocock, J. Bombay N.H. Soc. 55.- 278. Boothapundy, on 
the Ghats, north of Aramboly in Travancore, Southern India. 

Alacaca silenus group 
The name Silenus Goldfuss, 1820, is available for this species, which is well 
figured in Pocock, 1939, pi. 4, opposite p. 66, and is not likely to be confused with 
any other species. Tail length moderate. 

Macaca silenus Linnaeus, 1758 Lion-tailed Macaque 

Approximate distribution of species: Peninsular India; the Western Ghats, prin- 
cipally of Travancore and Cochin. 

Macaca silenus Linnaeus, 1 758 

1758. Simla silenus Linnaeus, Syst. Nat. loth ed. /.• 26. "Ceylon." 

1777. Cercopithecus veter Erxleben, Syst. Regn. An. 24. Not of Linnaeus, 1766. 

1792. Simla {Cercopithecus) veter alblbarbatus Kerr, Anim. Kingd. 64. 

1792. Slmia {Cercopithecus) silenus alblbarbatus Kerr, loc. cit. 

1793. Simla ferox Shaw, Mus. Leverian, 69. 
Range: as above. 

Alacaca nemestrina group 

Pocock would refer this to the subgenus Silenus if subgeneric division is required. 
It lacks the ruff of long greyish hair extending each side of face from temples to 
throat, which is a diagnostic character of AL silenus. Tail length medium. 

Macaca nemestrina Linnaeus, 1 766 Pig-tailed Macaque 

Approximate distribution of species: Assam, Burma, Siam, Malay States, Sumatra, 
Borneo, and a few small adjacent islands. 

(Macaca nemestrina nemestrina Linnaeus, 1766. Extralimital) 

■1766. Simla nemestrina Linnaeus, Syst. Nat. 12th ed. /.• 35. Sumatra. (Ranges north 
on the mainland about to Trang, Lower Siam.) 



Macaca nemestrina leonina Blyth, 1863 

1863. Macacus leoninus Blyth, Cat. Mamm. Mus. As. Soc. 7. Northern Arakaii, 

1869. Macacus andamanensis Bartlett, Land and ^Vater, 8: 57. Port Bhiir, Andaman 

Islands (introduced). 
iqo6. Macaca adusta Miller, Proc. U.S. Nat. Mus. 2g: 559. Champang, Tenasserim. 
iqo6. Macaca insulana Miller, Proc. U.S. Nat. Mus. sg: 560. Chance Island, Mergui 

1919. Macaca nemestrina indoc/uncnsis Kloss, J.N.H. Soc. Siam, j: 343. Lat Bua Kao, 

Eastern Siam. 
Range : Upper Burma to Tenasserim, Mergui Archipelago and Siam. 

Macaca nemestrina blythi Pocock, 1931 

1 93 1. Macaca nemestrina hlvlhii Pocock, J. Bombay N.H. Soc. j§: 305. Locality un- 
known. Described from a single captive specimen. Pocock says the distri- 
bution is unknown, "but probably some district of British India east of the 
'Ganges; ? Naga Hills, in Assam". 

Macaca irus group 
The subgeneric name Cynamolgiis Reichenbach, 1862, is available. Long-tailed 
species, differing from the sinica group in having the hair on the crown short. The 
differences between the two types are well figured in Pocock ( 1939, 35, 39, and pi. 5, 
opposite p. 79). 

Macaca irus Cuvier, 181 8 Crab-eating Macaque 

Appro.ximate distribution of species: Burma, Nicobar Islands, Indo-China, Siam, 
Malay States, Sumatra, Java, Borneo, and many small adjacent islands, east to 

(Macaca irus irus Cuvier, 1818. Extralimita!) 

1775. Simia cvnamolgus Schreber, Saugcth. /.• 91. N<it of Linnaeus, 1758. 
1 818. Macacus irus F. Cuvier, Mem. Mus. H.N. Paris, ^: 120. Sumatra (according to 
Chasen, 1940). Substitute for cvnamolgus Schreber. 

Macaca irus aurea Geoffroy, 1831 

1 83 1. Macacus aureus Geoffroy, Zool. Voy. de Belanger, 58, 76. Pegu, Burma. 

1 9 10. Pilhecus Vitus Elliot, Proc. U.S. Nat. Mus. ^8: 346. Domel Island, Mergui 

1915. Pithecus fascicularis Wroughton, J. Bombay N.H. Soc. 2j: 700. Not of Raffles, 

182 1." 
Range: Lower Burma, Tenasserim, Mergui .\rchipelago, South-Western Siam. 

Macaca irus umbrosa Miller, 1902 

1902. Macacus umhrusus Miller, Proc. U.S. Nat. Mus. 2^: 789. Little Nicobar Island, 

Bay of Bengal. Range; Great Nicobar, Little Nicobar and Katchal Island, 

Nicobar Islands. 



Macaca irus valida Elliot, 1909 

1909. Pithecus validus Elliot, Ann. Mag. N.H. 4: 252. Cochin-China. (Type skin in 

B.M. bearing label "Afacaca irus valida. The tail is imperfect, not complete 

as Elliot supposed.") 

Macaca irus atriceps Kloss, 191 9 

1919. Macaca irus atriceps Kloss, J. N.H. Soc. Siam, j: 347. Koh Kram Island, near 
Cape Liant, South-Eastern Siam. 

Macaca mulatta group 

Rhesus Lesson, 1840, is available if subgeneric division is required. Contains two 

closely allied species (mulatta and assamensis) which occur together, for characters 

see Pocock (1939, 33), and the Formosan M. crclopis seems to belong here. Tail of 

medium length and hairier than nemestrina; usually with no definite "cap" on crown. 

Macaca mulatta Zimmermann, 1780 Rhesus Macaque 

Approximate distribution of species: Kafiristan (Eastern Afghanistan), Kashmir 
Punjab, east to Nepal, Assam and Burma, south approximately to the Tapti River 
(Khandesh) and the Godavari in Northern Peninsular India; Siam, Indo-China- 
Szechuan and Yunnan, eastwards to Fukien and adjacent states in Southern China 
Hainan, Tibet, and the neighbourhood of Pekin, where perhaps introduced. 

Macaca mulatta mulatt.a Zimmermann, 1 780 

1780. Cercopithecus mulatta Zimmermann, Geogr. Gesch. Mensch. :?.■ 195. "India." 

1792. Simla (Cercopithecus) fulvus Kerr, Anim. Kingd. 73. "India." 

1 798. Simia rhesus Audebert, Hist. Nat. Singes, sig. i. Locality unknown. 

1800. Simia erythraea Shaw, Gen. Zool. /.• 33. Locality unknown. 

1840. Macaca (Pithex) oinops Hodgson, J. Asiat. Soc. Bengal, g: 12 12. Nepal Terai. 

1840. Macaca (Pithex) nipalensis Hodgson, J. Asiat. Soc. Bengal, 9.- 1212. Nepal Terai. 

1866. Imius sancti-johannis Swinhoe, P.Z.S. 556. North Lena Island, Hong Kong, 

China. For status, see G. Allen, 1938, Mamm. China & Mongolia, /.■ 284 
1868. Macacus lasiotus Gray, P.Z.S. 60, pi. 6. Szechuan, China. For status, see G. 

Allen, 1938, /.■ 284. 
1872. Macacus tchcliensis Milne-Edwards, Rech. Mamm. 227, pis. 32, 33. Mountains 

to the east of the Province of Tcheli (Chihli), North-Eastern China. For 

status, see G. Allen, 1938, /.• 284. 
1909. Pithecus littoralis Elliot, Ann. Mag. N.H. 4: 250. Kuatun, Fukien, South- 
Eastern China. 
1909. Pithecus brachyurus Elliot, Ann. Mag. N.H. 4: 251. Hainan. Not of H Smith 

1913. Pithecus brevicaudus Elliot, Rev. Primates, 2: 216, pi. 23. New name for brachyurus, 

1917. Macaca siamica Kloss J. N.H. Soc. Siam, 2: 247. Meping Rapids, below Chieng- 

mai, Siam. For status, see Pocock, 1939, Fauna Brit. India, Mamm. /.• 45. 
Range: Nepal, Bhutan, North Kamrup, Assam, Burma, Northern Peninsular India, 
Siam, Indo-China, Szechuan, Yunnan, to Fukien and adjacent states in Southern 
China, Chihli, Hainan. 



Macaca mulatta vestita Milne-Edwards, i.'ifjj 

1892. Miuaciis rcitilKS Milne-Edwards, Re\'. Gen. Seienccs, 671. Teiigri-nor, Tibet. 
G. .\llen lliinks this may be a synonym nl'the typical race. 

Mac.\ca mul.«lTta villosa .True, 1894 

1894. Macacus rhesus villosus True, Proc. U.S. Nat. Mus. ly: 2. Lolab, northern end 

of Wular Lake, about 40 miles north-west of Srinagar, Kashmir. Range ; 

Southern Kashmir, Upper Punjab, Kumaon, in Northern India. 

Macaca mulatta mcmahoni Pocock, 1932 

1932. Macaca mulatta mcmahoni Pocock, J. Bombay N.H. Soc. jj.- 544. Kootai, in 

Lower Chitrai, between the Bashgal Valley in Kafiristan and the Chitral 

Valley, 3,600 ft. Range: Kafiristan and Chitral. 

Macaca assamensis M'Clclland, 1839 Assainese Macaque 

Approximate distribution of species: Nepal, Sikkim, Bhutan, Assam, Northern 
Burma, south to the Sundarbans; Yunnan; Indo-Ghina. 

NL'^cACA assamensis assamensis M'Clelland 1839 

1839. Macacus assamensis M'Clelland, in Horsfield, P.Z.S. 148. Assam. 

1932. Macaca assamensis coolidgei Osgood, Field Mus. N.H. Zool. 18: 2og. Hoi Xuan, 

Annam, Indo-China. 
Range: Assam, Mishmi and Naga Hills, Northern Burma, Tonkin and Annam. 

Macaca assamensis pelops Hodgson, 1840 

1840. Macacus (Pithex) pelops Hodgson, J. Asiat. Soc. Bengal, g: 12 13. Nepal Kachar. 
1870. Macacus prohlcmaticus Gray, Cat. Monkeys, etc. B.M. 128. Dhalimkot, Bhutan. 
1872. Macacus rheso-similis Sclater, P.Z.S. 495, pi. 25. "East Indies." 

Range: Himalayas, from Mussoorie through Nepal and Sikkim, from 2,000 to about 
6,000 ft., to Bhutan. 

Macaca cyclopis Swinhoe, 1862 Formosan Macaque 

Approximate distribution of species: Formosa. 

Macaca cyclopis Swinhoe, 1862 

1862. Macacus cyclopis Swinhoe, P.Z.S. 350. Formosa. 

1863. Macacus {radiatus) affims Blyth, Gat. Mamm. Mus. As. Soc. 8. Formosa. 

Macaca speciosa group 
Lyssodes Gistel, 1848, is available for speciosa, a short-tailed monkey which 
dilTers from the other species in the abnormal external male genitalia (Pocock.) 
The Japanese M. fuscata resembles speciosa in its short tail, and in most other 
characters, but according to Pocock (1939, 70) differs from that species in the 
structure of the glans penis. 



Macaca speciosa F. Cuvier, 1825 Stump-tailed Macaque 

Approximate distribution of species : Szechuan and Yunnan, eastwards to Fukien 
and adjacent states in Southern China; Assam, Burma, Indo-China, south to 
Siamese Malaya. 

Macaca speciosa speciosa F. Cuvier, 1825 

1825. Macacus speciosus Cuvier, H.N. Mamm. 5, 47, Macaque a face rouge, 2. "East 

1 87 1. Macacus brunneus h.ndtKon, P.Z.S. 628. Kakhyen Hills, east of Bhamo, Yunnan- 

Burma border. M. brunneus=M. s. thibetanus, according to G. Allen. 
1912. Macacus (Magus) arctoides melli Matschie, S.B. Ges. Nat. Fr. Berlin, 308. West 

of Lochangho, Kwantung, Southern China. G. Allen uses this name for the 

South-Eastern Chinese form, but it is not distinguishable from brunneus, 

according to Pocock. 
191 2. Macacus [Magus) arctoides esau Matschie, loc. cit. 309. West of Lochangho, 

Kwantung, Southern China. 
1928. Pithecus pullus Howell, Proc. Biol. Soc. Washington, 41: 41. Near Kuatun, 

Fukien, Southern China. 
Range: Assam, Upper Burma, Southern China, Tonkin and Annam. 

Macaca speciosa arctoides Geoffroy, 1831 

1 83 1. Macacus arctoides I. Geoffroy, Zool. Voy. de Belanger, 61. Cochin-China. 
1854. Macacus ursinus Gervais, H.N. Mamm. /.• 93. Substitute for arctoides. Pro- 
visionally regarded as a valid race by Pocock. 

Macaca speciosa melanota Ogilby, 1839 

1839. Papio melanotus Ogilby, P.Z.S. 31. Type locality "said to be Madras". 

1872. Macacus rufescens Anderson, P.Z.S. 204. Singapore (where the animal does not 

occur, according to Chasen (1940), who lists it as a valid race from 

Peninsular Siam). 
1897. Macacus harmandi Trouessart, Le Naturaliste, //.■ 10. Chantabun, Southern 

Range: Tenasserim to Lower .Siam. 

Macaca speciosa thibetana Milne-Edwards, 1870 

1870. Macacus thibetanus Milne-Edwards, C.R. Acad. Sci. Paris, yo: 341. Near 

Moupin, Szechuan, China. Emended to Macacus tibetanus Milne-Edwards, 

1872, Rech. Mamm. 244, pis. 34, 35. 

Macaca fuscata Blyth, 1875 Japanese Macaque 

Approximate distribution of species: Japan; including Shikoku and Kiushiu, 
Hondo and Yakushima. 

Macaca fuscata fuscata Blyth, 1875 

1875. Macacus fuscatus Blyth, J. Asiat. Soc. Bengal, 44. (extra number). Cat. Mamm. 

& Birds, Burma, 6. Japan. 
1842. Itiuus speciosus Temminck, Fauna Japonica, 9. Not of Cuvier, 1825. 
1909. Inuus speciosus japanensis^c\\'wcyeT,A.nihroY>.-'Zoo\.\]nteTi\ich. Miinchen, 1-192. 



Macaca fuscata YAK.UI Kuroda, 1941 

1941. Macaca fuscata yakui Kuroda, Monogr. Jap. Mamm. 273. Yakushima Island, 

Macaca sylrana group 
(= Macaca sensu stricto. Fur characters, see above, page 194) 

Macaca sylvana Linnaeus, 1758 Barbary Ape 

Approximate distribution of species: Morocco and Algeria. (Introduced in 

Mac.\c.\ sylvana Linnaeus, 1758 

1758. Simia sylvamis Linnaeus, Syst. Nat. loth ed. /.■ 25. ("In Africa, Ceylona.") 

Barbary Coast. 
1766. Simia irmus Linnaeus, Syst. Xat. 12th ed. /.• 35. "Africa." 
1799. Simia pithecus Schreber, Saugeth. Suppl. /.• pi. 4b. 
1812. Imiia ecaudatus E. Geoffroy, Ann. Mus. H.X. Paris, 79.- 100. Mediterranean 

coast of Africa and Gibraltar. 
1B63. Pithecus pygmaetii Reichenbach, \'ollstand. Xat. .\nen, 145. 
Range: as above. 

Genus PAPIO Muller, 1773 

1773. Papio Muller, \"ollstand. Natursyst. /: iiH. Usually applied to the baboons 
(except the hamadryas and gelada), but according to Hopwood the type of 
this genus should be taken as Simia sphinx Linnaeus (the West African 

1795. Cnwcephalus Cuvier & Geoffroy, Mag. Encyclop. j; 462. Simia cynocephalus 
Linnaeus. Xot of Boddaert, 1768. 

1824. Mandrillus Ritgen, Xat. Eintheil. Saugeth. 33 (Tafel). (teste Palmer.) Simia 
maimon Linnaeus and Simia mormon Alstromer, both of which are synonyms 
of Simia sphinx Linnaeus, according to G. Allen. 

1830. Chacropithecus Gervais, Diet. Pittor. Hist. Xat. 8: 90 (prior to 11 May). Simia 
cynocephalus Linnaeus. Valid as a subgenus. If Papio is used for the mandrills, 
then Chacropithecus becomes the name for the baboons (except the hama- 
dryas and the gelada j. 

1839. Chacropithecus Blain\ille, Osteogr. Mamm. Pithecus, 39 ' 14 June). Simia cyno- 

cephalus Linnaeus. 

1840. Hamadryas Lesson, Spec. Mamin. 107. X(jt ofHubner, 180G. Hamadryas choero- 

pithecus Lesson = Simia hamadryas Linnaeus. 

1862. Choiropithecus Reichenbach, \'ollstand. Xat. Affen, 151. Simia porcaria Boddaert. 

1925. Comopithccus ]. Allen, Bull. Amer. Mus. X.H. ^y: 312. Simia hamadryas Lin- 
naeus. To replace Hamadryas Lesson, preoccupied. Valid as a subgenus. 

I species in Asia : 

Papio hamadryas, page 201 


Only one species of this genus occurs in Asia, the others being confined to Ethiopian 
Africa. This species is sometimes separated generically as Comopithecus, e.g. by G. 
Allen and Simpson. On the other hand, even an extreme splitter like Elliot referred 
all Baboons to one genus, Papio. Hopwood, 1947, P-Z-^- ^'7' 533~536, has shown 
that the type oi Papio is P. sphinx, the Mandrill, currently referred to a distinct genus 
Mandrillus, and he would call the other Baboons of Africa Choeropithecus Blainville, 
which is antedated by Choeropithecus Gervais. However, we suggest subgeneric rank for 
all three groups. 

The copious mane on the head and shoulders of the male seems to be the most 
obvious distinguishing character of the subgenus Comopithecus. 

Subgenus COMOPITHECUS J. Allen, 1925 

Fapio hamadryas Linnaeus, 1758 Sacred Baboon 

Approximate distribution of species: Arabia; Somaliland, Abyssinia, Sudan. 

(Papio hamadryas hamadryas Linnaeus, 1758. Extralimital) 

1758. Simia hamadryas Linnaeus, Syst. Nat. loth ed. /.• 27. Egypt (where now extinct). 
1758. Simia cynamolgos Linnaeus, Syst. Nat. loth ed. /.■ 28. Upper Egypt. 
1840. Hamadryas chaeropithecus Lesson, Spec. Mamm. 109. Abyssinia, Arabia, Egypt. 
1870. Hamadryas aegyptiaca Gray, Cat. Monkeys, etc. B.M. 34. New name for Aama- 

dryas Linnaeus. 
Range: Eastern Ethiopia and Eastern Sudan, mainly in the lowlands. 

Papio hamadryas .'Vrabicus Thomas, 1900 

1900. Papio arabicus Thomas, P.Z.S. i8gg: 929; and igoo: 96. Subaihi country, 
about 60 miles north-west of Aden, Southern Arabia. 

Subfamily C o 1 o b i n a e 

Genus RfflNOPITHECUS Milne-Edwards, 1872 

1872. Rhinopithecus Milne-Edwards, Rech. H.N. Mamm. 233. Semnopithecus roxellana 

1924. Presbytiscus Pocock, Abstr. P.Z.S. 17. Rhinopithecus avunculus Dollman. Valid as 

a subgenus. 

Pocock seems to base his name Presbytiscus chiefly on the fact that the digits of the 
hand and feet are relatively longer than in Rhinopithecus. The name Presbytiscus is 
ignored by Simpson (1945). 

The other members of the genus seem to be not very well known. G. Allen (1939, 
300) follows Elliot in listing the three named forms as distinct species. It is difficult to 
believe that three forms, not occurring together (see Allen's distribution map) and 
differing apparently only in details of colouring (which might even be seasonal) are 


good species, and until the contrary is proved we prefer to regard them as repre- 
sentatives of one species, for which roxellanae is the first name. 

2 species : Rhino/iit/nriis avunculus, page 202 
Rhiiiopitlurns roxellanae, page 202 

Subgenus RHINOPITHF.CUS Milne-Edwards, 1872 

Rhinopithecus roxellanae Milne-Edwards, 1870 Snub-nosed Monkey 

Grilden Monkey 

Approximate distribution of species, as here understood: China, states of Szechuan 
,into Southern Kansu), Yunnan and Kweichow. 

Rhinopithecus roxellaxaf, roxellanae Milne-Edwards, 1870 

1870. Scmnopilhecus roxellana Milne-Edwards, C.R. Acad. Sci. Paris, yo: 341. Near 

Moupin, Szechuan, C^hina. 
1872. Semnopithecus roxellanae Milne-Edwards, Rech. H.N. Mamm. 233-243, pis. 

3*^, 37- 

Rhinopithecus roxellanae bieti Milne-Edwards, 1897 

1897. Rhinopithecus bieli Milne-Edwards, Bull. Mus. H.N. Paris, 3: 157. Kiape, a day's 
journey from Atuntze (left bank Mekong River), North- Western Yunnan, 
China. See also Milne-Edwards & Pousargues, 1898, Nouv. Arch. Mus. 
H.N. Paris (3), 10: 121-142, pis. 9-12. 

Rhinopithecus roxell.-\nae brelichi Thomas, 1903 

1903. Rhinopithecus brelichi Thomas, P.Z.S. 224, pi. 21. Probably from Northern 
Kweichow (? Van Gin Shang Range, 29"' N., 108° E.), China. 

Subgenus PRESBYTISCUS Pocock, 1924 

Rhinopithecus avunculus Dollman, 19 12 Tonkin Snub-nosed Monkey 

Approximate distribution rif species: Tonkin, in Indo-China. 

RHiNOPiriiEcus .wuNCULUs Dollman, 191 2 

1912. Rhinopithecus avunculus Dollman, Abstr. P.Z.S. 18; P.Z.S. 503. Yen Bay, Song- 
koi Ri\er, Tonkin, Indo-China. 

Genus PYGATHRIX E. Geoffroy, 18 12 

1812. P}xathrix GeoflVoy, Ann. Mus. H.X. Paris, ic^: 90. Simla ncmaeus Linnaeus. 

It should be noted that although the International Commission of Zoological 
Nomenclature, in Opinion 114, suppressed the name Pithecus (1795, Cuvier & 
Geoffrov, Mas,. Encvct. •5.- 462, based on the unidentifiable Simla veter Linnaeus), 


Chasen (1940) declared himself a rebel and continued to use it. Allen, 1938, Mammals 
of Mongolia & China, also continued to use the name. Allen, unlike Chasen, did not 
attempt to explain his rejection of the Commission's authority and it is interesting to 
note that one year later, in his Checklist of African Mammals, he quoted Opinion 1 14, 
without protest, as the authority for the suppression of Simla. At all events, so far as 
we are concerned, and we believe that most mammalogists are with us, Pithecus is 
dead. Therefore, if all the Langurs are regarded as being congeneric, Pygathrix is the 
valid name. 

Pocock (1939) refers the Indian langurs to four genera: Presbytis, Trachy pithecus, 
Kasi and Semnopithecus — for reasons which do not convince us, and we here follow 
Thomas, Simpson and Osgood in dividing the langurs into two genera: Pygathrix for 
the species nemaeus, and Presbytis for the remainder. 

I species: Pygathrix nemaeus, page 203 

Pygathrix nemaeus Linnaeus, 1771 Douc Langur 

Approximate distribution of species: Indo-China (Annam, Laos, Cochin-China), 
and has been recorded from Hainan. 

For characters and revision, see Pocock, 1935, P.^.S. igj^: 958. 

Pygathrix nemaeus nemaeus Linnaeus, 1771 

1 77 1. Simla nemaeus Linnaeus, Mant. Plant, 521. Cochin-China. 

Pygathrix nemaeus nigripes Milne-Edwards, 1871 

1871. Semnopithecus nigripes Milne-Edwards, Bull. Nouv. Arch. Mus. H.\. Paris, 6: 

7, pi. I. Saigon, Cochin-China. 
1926. Presbytis nemaeus mol Kloss, Ann. Mag. N.H. 18: 214. Langbian Peak, 5,500- 

6,500 ft., Southern Annam, Indo-China. 

Genus PRESBYTIS Eschscholtz, 182 1 

1 82 1. Presbytis Eschscholtz, in Kotzebue Reise, j: 196, pi. Presbytis mltratus Esch- 

scholtz = Simla aygula Linnaeus, the Sunda Islands Leaf Monkey, from 

1822. Semnopithecus Desmarest, Mamm. 2: 532. Simla entellus Dufresne. 

1862. Trachyplthecus Reichenbach, Vollstand. Nat. Affen, 89. Semnopithecus pyrrhus 

Horsfield, from Java. 
1862. Kasl Reichenbach, Vollstand. Nat. Affen, loi. Cercoplthecus johnll Fischer. 
1879. Coryplthecus Trouessart, Rev. Mag. Zool. (3), j: 53. Semnopithecus frontatus 

Muller, from Borneo. 
1879. Lophoplthecus Trouessart, Rev. Mag. Zool. (3), j: 53. Semnopithecus rublcundus 

Muller, from Borneo. 
1879. Presbypithecus Trouessart, Rev. Mag. Zool. (3), 7.- 56. Substitute for Presbytis 

Reichenbach, 1862, not of Eschscholtz, 1821. 

o 203 

palaearc'.tk; and Indian mammals 1758-1946 

9 species in the area co\ered by this list : 

Presbytis aislalus, page 208 Presbytis ohscurui, page 209 

Presbytis entellus, page 204 Presbytis phayrei, page 209 

Presbytis fran(oisi, page 210 Presbytis pileatiis, page 208 

Presbytis johni, page 207 Presbytis senex, page 206 
Presbytis melalophos, ,pvige 207 

\Vc do not know why Chasen ( 1 940) Hsted a long group of races as forms of 
Jemoralis which dates from 1838, including among them melalophos, which dates from 
1 82 1 thus clearly taking priority ; nor why he lists cristalus, which dates from 1 82 1 , as 
a subspecies of pyrrhus, which dates from 1823. He has dealt similarly with Sus 
crislatus 1839 (making vittatus 1828 a subspecies), and Rattus rapit 1903 (making 
lepturus 1879 a subspecies), and is likely to be widely followed. 

See Pocock, 1935, P-ZS- 1934: 895, for a review of the species to the east of the 
Bay of Bengal, and 1939, Fauna Brit. India, Mamm. /, for the species inhabiting India. 

Pocock restricted the name Presbytis to the ayoula group, and he recognized nine 
species, including P. aygula Linnaeus, 1758 (from Java, Sumatra and Borneo), P. 
melalophos and P.femoralis. Chasen (1940) only recognizes four species in this group, 
merging melalophos and femoralis (as mentioned above). We tentatively follow Chasen 
in his classification, although we are not sure that melalophos as here understood is 
clearly definable. Pocock referred the \Vestern Indian species entellus to the genus 
Semnopithecus, and the species senex and johni to the genus Kasi ; distinguishing characters 
for these groups will be found in his work on the mammals of India. P. johni is closely 
allied to senex, and could be regarded as a very distinct subspecies of it. The remaining 
five species now under discussion were referred by Pocock to the genus Trachypithecus. 
P. franpisi seems much the most distinct of these, characterized by black colour com- 
bined with very sharply contrasted white head, or cheeks, or rump. In this it resembles 
the extralimital P. poten-iani, from which it differs by some skull characters. The 
remainder arc very closely allied to each other, but three of them occur together in 
Burma, and Pocock has given characters by which apparently they may be dis- 
tinguished. It must be noted that cristatus is the prior name for this section of the 

Presbytis entellus group 
= the genus Semnopithecus ^Desmarest, 1822) of Pocock, 1939. 

Presbytis entellus Dufresne, 1797 Langur (Entellus Monkey) 

Approximate distribution of species: Ceylon, Peninsular India, northwards to 
Sikkim and Kashmir, and extreme Southern Tibet. 

Presbytis entellus entellus Dufresne, 1797 

1797. Simia entellus Dufresne, Bull. Soc. Philom. Paris, /, 7: 49. Bengal, Ind 
Range : Bengal to Gujerat and Kathiawar. 




Presbytis en'tellus schistaceus Hodgson, 1840 

1840. Semnopithecus schistaceus Hodgson, J. Asiat. Soc. Bengal, g: 1212. Nepal Terai. 
(Not schistaceus Blanford, 1891.) 

1840. Semnopithecus nipalensis Hodgson, J. Asiat. Soc. Bengal, g: 1212. 

1928. Pithecus entellus hector Pocock, J. Bombay N.H. Soc. ^2: 481. Sitabani, Ramna- 

gar, Kumaon, 2,000 ft., Northern India. 
Range: Nepal Terai, Oudh, Kumaon, Garwhal. 

Presbytis entellus hypoleucos Blyth, 1841 

1 84 1. Semnopithecus hypoleucos Blyth, J. Asiat. Soc. Bengal, 10: 839. Travancore, 

Southern India. 

Presbytis entellus dussumieri Geoffroy, 1843 

1843. Semnopithecus dussumieri I. Geoffroy, C.R. Acad. Sci. Paris, i^: 719. Malabar 

coast, India. 

Presbytis entellus anchises Blyth, 1844 

1844. Presbytis anchises Blyth, J. Asiat. Soc. Bengal, 75.- 470. Deccan, India. Range: 

Central Provinces and Eastern Ghats. 

Presbytis entellus priam Blyth, 1844 

1844. Semnopithecus pallipes Blyth, Ann. Mag. N.H. 13: 312 (April). (See Pocock, 

1939, /.• 109, footnote, on synonymy.) 
1844. Semnopithecus priam Blyth, J. Asiat. Soc. Bengal, i^: 470 (October). Coro- 

mandel coast, India. 
1847. Semnopithecus priamus Blyth, J. Asiat. Soc. Bengal, 16: 732. 
Range: the Dharmapuri, Shevaroy and Palkonda Hills, and Nilgiri Hills, India. 

Presbytis entellus thersites Blyth, 1847 

1847. Presbytis thersites Blyth, J. Asiat. Soc. Bengal, 16: 1271. Trincomalee, Ceylon. 
Range: Ceylon and apparently Travancore (Pocock). 

Presbytis (?) entellus lania Elliot, 1909 

igog. Presbytis lania Elliot, Ann. Mag. N.H. 4: 273. Chumbi Valley, extreme 
Southern Tibet. 

Presbytis entellus achilles Pocock, ig28 

ig28. Pithecus entellus achilles Pocock, J. Bombay N.H. Soc. 32: 478. Satthar Hill, 

Gorkha, 12,000 ft., 50 miles north-west of Katmandu, Nepal. 
1888. Semnopithecus schistaceus Blanford, Mamm. Brit. India, 30, not of Hodgson, 

Range: Sikkim and Nepal, at high altitudes; ? Kashmir. 

Presbytis entellus ajax Pocock, ig28 

1928. Pithecus entellus ajax Pocock, J. Bombay N.H. Soc. ;^2: 480, pi. 2, fig. i. 
Deolah, in Chamba, 6,000 ft., Punjab. Range: Chamba, Kangra and Kulu, 
at high altitudes; ? Kashmir. 



Presbytis entellus achates Pocock, 1928 

1928. Pithecus (ntdlus achates Pocock, J. Bombay N.H. Soc. ji'.- 488. Haunsbhavi, 
Dharvvar, 2,000 ft., India. Ranejc: Dhanvar, Bellary and Kanara. 

Presbytis entellus iiilus Pocock, 1928 

1928. Pithecus entellus iulus Pocock, J. Bombay N.H. Soc. jp; ^go. Jog, Gersoppa 
Falls, on Kanara-Mysore border, 1,300 ft., India. 

Presbytis entellus aene.'vs Pocock, 1928 

1928. Pithecus entellus aencas Pocock, J. Bombay N.H. Soc. ;^2: 492. Makut, Southern 

Coorg, 250 ft., India. Range: Southern Coorg, from Makut to Wottekolli, 

2,000 ft. 

Presbytis entellus eliss.\ Pocock, 1928 

1928. Pithecus entellus elissa Pocock, J. Bombay N.H. Soc. 32: 493. Nagarhole, South- 
Eastern Coorg, India. 

Presbytis entellus pri.\mellus Pocock, 1928 

1928. Pithecus entellus priamellus Pocock, J. Bombay N.H. Soc. jj:?.' 494. Shernelly, 
Cochin, India. 

Presbytis senex group 
= the genus K'asi (Reichcnbach, 1862) of Pocock, 1939. 

Presbytis senex Erxleben, 1777 Purple-faced Langur 

Approximate distribution of species: Ceylon. 

Presbytis senex senex Erxleben, 1777 

1777. Cercopithecus senex Erxleben, Regn. Anim. 24. "Hills of Southern Ceylon." 

1852. Preshrtis alhinus Kelaart, Prodr. Faun. Zeyl. 7. Matalc, Central Province, 

1927. Pithecus philbricki VhiWips, Ceyl.J. Sci. Sec. B, /./.• 57. Kaiitalai, East Province, 

200 ft., near Trincomalee, Ceylon. 
Range: "The hills east of Matale and Madulkellc up to 5,000 ft., also the low- 
country dry zone of the N.C.P., N.W.P., E.P. and C.P., Ceylon." 

Presbytis senex vetulus Erxleben, 1777 

1777. Cercopithecus vetulus Erxleben, Rcgn. Anim. 25. Ceylon. 

1780. Cercojiitliecus kephaloplerus Zimmcrmann, Geogr. Ges. j?.- 185. "cephalopterus" of 

many subsequent authors. Ceylon. 
Range; wettest parts of lowlands of Western and South- Western Ceylon. 

Presbytis senex nestor Bennett, 1833 

1833. Sernnopithecus nestor Bennett, P.Z.S. 67. Ceylon, probably Rayigam. 

1923. Pithecus vetulus phillipu Hinton, ,\nn. Mag. N.H. //.• 510. Gonapola, Panadura 

district, Ceylon. 
Range: lnw-country wet zone of Western Province, Ceylon. 



Presbytis senex monticola Kelaart, 1850 

1850. Presbytis cephalopterus var. monticola Kelaart, J. Ceylon Br. Asiat. Soc. 2: 207 

(321 of 1887, reprint). Nuwara Eliya, Ceylon. 

1851. Presbytis ursinus Blyth, J. Asiat. Soc. Bengal, 20: 155. Nuwara Eliya, Ceylon. 
Range : hill ranges of Ceylon, above 4,000 ft. 

Presbytis johni Fischer, 1829 John's Langur 

Approximate distribution of species: Coorg, Nilgiri and Palni Hills, in Southern 

Presbytis johni Fischer, 1829 

1829. Cercopithecus johnii Fischer, Syn. Mamm. 25. Tellicherry, Southern India. 

1834. Semnopithecus cucullatus I. Geoffroy, Zool. Voy. Belanger, 38, pi. i. The Ghats, 

1840. Semnopithecus jubatus Wagner, Schreber Saugeth. Suppl. /.• 305. Southern India. 
Range; Southern India; Western Ghats, from Coorg southwards, Nilgiri, Anamalai, 

Brahmagiri, Tinnevelly and Palni Hills, usually not below 3,000 ft. (Pocock). 

Presbytis aygula group 
= part of the genus Presbytis as restricted by Pocock, 1939. 

Presbytis tnelalophos Raffles, 182 1 Banded Leaf Monkey 

Approximate distribution of species: Tenasserim, Siam, Malay States, Sumatra, 
Borneo, and some adjacent small islands. 

(Presbytis melalophos melalophos Raffles, 1821. Extralimital) 
182 1. Simia melalophos Raffles, Trans. Linn. Soc. London, 73.- 245. Bencoolen, 

(Presbytis femoralis Martin, 1838, Charlesworth's Mag. N.H. 2: 436, Singapore, 
is also extralimital. Pocock refers the race which occurs in Tenasserim to 

(It should be noted that the form Semnopithecus siamensis Mtiller & Schlegel, 1841, 
Verh. Nat. Ges. Ned. Overz. Bezitt. ^ool. Mamm. 60, listed by Elliot with several 
synonyms, is a race of melalophos but came from the Malay States, not from Siam, 
and so is extralimital.) 

Presbytis melalophos robinsoni Thomas, 19 10 

1910. Presbytis robinsoni Thomas, Abstr. P.Z.S. 25. P.Z.S. 635. Ko-khau, Trang, 

Lower Siam. Based, according to Pocock, on a partial albino, but ante- 
dating the next, which Pocock adopted. 

191 1. Presbytis neglecta keatii Robinson & Kloss, J. Fed. Malay States Mus. 4: 174. 

Ko-khau, Trang, Lower Siam. For status see Chasen, 1940, Handlist 
Malaysian Mamm. 74. 
Range: North Malay Peninsula, Junk Seylon Island, Tenasserim, and west of 
Bangkok, in Siam. 


palakarctr: and Indian mammals 1758-1946 

Prcshytis crislatiis group 
= the genus Trachypithecus fReichenbach, 1862) of Pocock, 1939. 

■Presbytis cristatus Rallies, 1821 Silvered Leaf Monkey 

Approximate distriijution of species: Tenasserim, Siam, Indo-China, Malay 
States, Sumatra, Java, Borneo, and various small adjacent islands. 

(Presbytis cristatus cristati's Raffles, 1821. Extralimital) 

1 82 1. Simla cristata Raffles, Trans. Linn. Soc. London, /jj.- 244. Bencoolen, Sumatra. 

(This antedates Semnopitheciis pyrrhus Horsficld, 1823, ^ool. Res. Java, pt. 7 
(unpaged), pi. 3, Java. For date of publication, see Matthews, 1919, Birds of 
Australia, y, 5: 475, and Oberholser, 1921, Proc. Biol. Soc. Washington, j^: 163- 

Presbytis cristatus germaini Milne-Edwards, 1876 

1876. Scmnopithecus germani (misprint tor germaini) Milne-Edwards, Bull. Soc. Fhilom. 

(6), //.• 8. (The collector's name was Germain, and most authors have 

emended to germaini.) Cochin-China and Cambodia. 
1909. Presbytis margarita Elliot, Ann. Mag. N.H. ./; 271. Langbian, Annam. 
1916. Presbytis germaini mandihularis Kloss, P.Z.S. 32. Koh Chang (Island), South- 

Eastern Siam. 
1919. Presbytis cristatus koratensis Kloss, J. N.H. Soc. Siam, j.- 340. Lat Bua Kao, 30 

miles west of Korat, .Siam. 
Range: Indo-China and Siam. 

Presbytis crist.\tus atrior Pocock, 1928 

1928. Pi thecus pyrrhus atrior Pocock, J. Bombay N.H. Soc. ^2: 673. Ye Forest, 500 ft., 
south of -Moulmcin, in .\taran district of Tenasserim. Range includes South- 
Western .Siam. 

(?) 1863. Presbytis barbel Blyth, Cat. Mamm. .Mus. .A.siat. Soc. 14. Tipperah Hills. 
Not barbel Blyth, 1847. 

Presbytis pileatus Blyth, 1843 Capped Monkey 

.\pproximate distribution of species: Assam and Burma. ? Yunnan (Pocock, 
1939, '3' (footnote), suggests that G. Allen's Plthecus ohscurus luirbei may be this 

Presbytis pileatus pileatus Blyth, 1843 

1843. Semnoplthecus pileatus Blyth, J. Asiat. Soc. Bengal, /-■.• 174. Locality unknown. 

"received from Barrackpore, stated to be Malayan" (Blyth); "no doubt 

Assam" (Pocock). 
1851. Semnoplthecus argentatus Horsfield, Cat. Mamm. E. India Co. 7. Sylhct. 
Range: Garo, Khasi, Jaintia and .\aga Hills. Assam, above 4,000 ft. 



Presbytis pileatus shortridgei Wroughton, 1915 

1915. Presbytis shortridgei Wroughton, J. Bombay N.H. Soc. 24: 56. Homalin, Upper 
Chindwin, Burma. 

1915. Presbytis shortridgei belliger Wroughton, J. Bombay N.H. Soc. 24: 57. Hkamti, 

Upper Chindwin, Burma. 
Range: eastern side of Upper Chindwin, Burma. 

Presbytis pileatus brahma Wroughton, 1916 

1916. Presbytis brahma Wroughton, J. Bombay N.H. Soc. 24: 654. Seajulia, Dafla 

Hills, Northern Lakhimpur, Upper Assam. 

Presbytis pileatus durg.a Wroughton, 1916 

1916. Presbytis durga ^Vroughton, J. Bombay N.H. Soc. 24: 655. Cachar, Assam. 

1923. Pithecus pileatus saturatus Hinton, J. Bombay N.H. Sor. 2g: 81. Bara Hapjan, 

Lakhimpur, Upper Assam. 
Range: Lakhimpur, in Upper Assam, south to Naga Hills, Cachar, Tipperah, Chitta- 

gong, and western side Upper Chindwin, Burma. (In Assam, occurring at 

lower levels than the typical race.) 

Presbytis pileatus tenebricus Hinton, 1923 

1923. Pithecus pileatus tenebricus Hinton, J. Bombay N.H. Soc. 2g: 81. Matunga River, 
Northern Kamrup. Range includes Assam, north of the Brahmaputra. 

Presbytis obscurus Reid, 1837 Dusky Leaf Monkey 

Approximate distribution of species: Tenasserim, Lower Siam, Malay States, and 
some small adjacent islands. 

(Presbytis obscurus obscurus Reid, 1837. Extralimital) 

1837. Semnopithecus obscurus Reid, P.Z.S. 14. Malacca, see Chasen (1940). 

Presbytis obscurus sanctorum Elliot, 19 10 

1910. Pygathrix sanctorum Elliot, Proc. U.S. Nat. M.w. 38: 351. St. Matthew Island, 
Mergui Archipelago. 

Presbytis obscurus flavicauda Elliot, 19 10 

1 910. Pygathrix flavicauda Elliot, Proc. U.S. Nat. Mus. 38: 352. Trang, Lower Siam. 

1916. Presbytis obscura smithi Kloss, J.N.H. Soc. Siam, 2: 5. Klong Bang Lai, Patiyu, 

Peninsular Siam. 
1935- Trachypithecus obscurus corax Pocock, P.Z.S. 1934: 944. Tenasserim Town, 

Range: from Northern Malaya northwards to Tavoy, in Tenasserim, and to Pech- 

buri district, South-\Vestern Siam. 

Presbytis phayrei Blyth, 1847 Phayre's Leaf Monkey 

Approximate distribution of species: Burma, north to Bhamo, Tenasserim, Siam, 
? Indo-China. 

Pocock (1928) regarded these forms as further races oi obscurus, but in his later 
work kept them apart on the ground of their simultaneous occurrence in Tenasserim. 


PALAKARCTK; and INDIAX mammals 17^,8-1946 

Presbytis phavrei phavrei Blyth, 1847 

1847. Presbytis phayrei Blyth, J. Asiat. Soc. Bengal, 16: 733. Arakan, Burma. 

1847. Presbytis barbei Blyth, J. Asiat. Soc. Bengal, 16: 734. Tipperah Hills. (See 
Pocock, 1939, Fauna Brit. India, Mamm. /.• 130-131, for notes on synonymy 
of this form. Not barbei Blyth, 1863, and evidently not P. obseuriis barbei of 
G. Allen, 1938, Mamm. China & T^Iongolia, /.• 294, which Pocock suggests 
might be a form of P. pileatus.) 

1909. Presbytis melamera Elliot, .\nn. Mag. N.H. ./.■ 'if)"]. Cadu Cliaung, near Bhamo, 
North-Eastcrn Burma. 

Range: Burma, as far north as Bhamo, south to Pegu. 

Presbytis phayrei crepusculus Elliot, 1909 

1909. Presbytis crepusnila Elliot, Ann. Mag. N.H. _/.• 271. Mt. Mulaiyit, 5,000 ft., 

1909. Presbytis erefniseula wrouohtoni Elliot, Ann. Mag. N.H. 4: 272. Pachebon, Clcntral 

1919. Presbytis art^rnteiis Kloss, J.N.H. Soc. Siam, j: 338. Lat Bua Kao, west of Korat, 

Ranges to Laos and Annam? 

Presbytis phayrei shanicus Wroughton, 191 7 

191 7. Pitheeus shanicus Wroughton, J. Bombay N.H. Soc. 2§: 47. Se'en, Hsipaw State, 

Shan States, Burma. Range: North Shan States and their neighbourhood to 

east of Irrawaddy, in dry zone (if Burma. 

Presbytis (?) phayrei ruhei Knottnerus-Meyer, 1933 

1933. Presbytis ruhei Knottnerus-Meyer, Zool. Garten, Leipzig, 6: 259. Sangora, 

Southern .Siam. From description may belong in this species, but status not 


Presbytis fran^oisi Pousargues, 1898 Francois' Monkey 

Approximate distribution of species: Kwangsi, in S(3uthern China; and Tonkin, 
Laos and Annam, Indo-China. 

For status of this species and for some skull characters by which the species or 
group seems distinguishable, see Pocock, 1935, P.^.S. 1934: 956-958. Pocock recog- 
nized four species, and so did Osgood (1932) who gave a key to them. But as they do 
not appear to occur together, and the region is a small one, we propose provisionally 
to regard them as races of the same species. 

Presbytis francoisi francoisi Pousargues, 1898 

1898. Semnopithecus francoisi Pousargues, Bull. Mus. H.N. Paris, 4: 319. Lungchow, 
Province of Kwangsi, Southern China. Range: southwards into Tonkin. 

Presbytis (?) fra.n'COisi poliocephalus Trouessart, 191 1 

191 I. Semriopithecui { Lophopithecus) poliocephalus Trouessart, Ann. NLig. N.H. S: 271. 
K.ii-Chin, Ncirth-Eastern Tonkin, Lido-China. 



Presbytis (?) FRAN901S1 LAOTUM Thomas, 1921 

1 92 1. Pithecus laotum Thomas, Ann. Mag. N.H. y: 181. Ban Na Sao, Mekong River, 
17.30° N., Laos, Indo-China. 

Presbytis (?) francoisi delacouri Osgood, 1932 

1932. Pithecus delacouri Osgood, Field Mus. N.H. Zool. 18: 205. Hoi Xuan, North- 
Eastern Annam, Indo-China. 

The name Simia veter Linnaeus, 1766, Syst. Nat. ed. 12, /.■ 36, supposed to have 
come from Ceylon, is held to be unidentifiable. 


Subfamily Hylobatinae 

This subfamily is given family rank by some authors. 
Genus : Hylobates, page 2 1 1 

Genus HYLOBATES Illiger, 181 1 

181 1. Hylobates Illiger, Prodr. Syst. Mamm. 67. Homo lar Linnaeus. 
1841. Symphalangus Gloger, Gemeinn. Naturg. /.- 34. Symphalangus syndactylus Gloger 
= Simia syndactylus Raffles. Valid as a subgenus. 

1932. Brachitanytes Schultz, J. Mamm. 13: 369. Symphalangus klossii Miller, from 

South Pagi Island, west of Sumatra. 

1933. Nomascus Miller, J. Mamm. i^: 159. Hylobates leucogenys Ogilby. Valid as a 


On the characters of the subgenera Hylobates, see Miller, 1933, J. Mamm. /^.- 158, 

4 species in the area covered by this list: 

Hylobates concolor, page 2 1 2 
Hylobates hoolock, page 2 1 2 
Hylobates lar, page 2 1 2 
Hylobates syndactylus, page 2 1 3 

Authors are not in agreement as to the full number of species in this genus, but the 
above four are universally admitted. See Pocock, 1927, P.^-S. 719, The Gibbons of 
the Genus Hylobates. Also Chasen, 1940, Handlist Malaysian Mammals, 63, in which 
certain forms referred to H. lar by Pocock are given specific rank ; one of these, agilis, 
occurs with lar in the Malay States. 


PALAEARtrnC: and INDIAN MAMMALS 1758-1946 
Subgenus HI'LOBATES Illiger, 181 1 

Hylobates lar Linnaeus, 1771 Lar Gibbon 

Approximate distribution ofspecies: Sumatra, Malay States, South-Western Siam, 
Tenasserim, Southern Indo-China (Cambodia). 

(Hylobates lar Linnaeus, 1771. ExtraHmital) 
I 77 1. Homo lar Linnaeus, NLint. Plant, 521. Malacca. 

Hylobates lar entelloides L Geoffrey, 1842 

1842. Hvlobatfi entelloides I. Gcoffroy, C.R. Acad. Sci. Paris, 75.- 717. Malay Penin- 
sula, about latitude 12° N. Range: Lower Siam, Tenasserim. 

Hylob.\tes lar pileatls Gray, 1861 

1861. Hvlohales fnleatus Gray, P.Z.S. 136. Cambodia. Range includes South-Eastern 

Hylobates hoolock Harl.m, 1834 Hoolock Gibbon 

Approximate distribution ofspecies: Yunnan, Assam and Burma. 

Hylobates hoolock Harlan, 1834 

1834. Simia hoolock Harlan, Trans. Amer. Phil. Soc. ^: 52, pi. 2. Garo Hills, Assam. 
1834. Hylobates fiiscus Winslow Lewis, J.N.H. .Soc. Boston, /, i : 40, pis. i and 2. 

"Vicinity of Himalaya Mountains." 
1837. Hylobates choromandus Ogilby, P.Z.S. 6g. Locality unknown. 
1840. Hylobates scyritus Ogilby, Royle's lUustr. Bot. Himal., Ix. Assam. 
Range: Assam, Cachar and Chittagong, through Upper Burma, to north Shan States 
and Western Yunnan. 

Subgenus NOMASCUS Miller, 1933 

Hylobates concolor Harlan, 1826 Black Gibbon 

Approximate distribution ofspecies: Hainan, Lido-China, Siam. 

Hylobates concolor concolor Harlan, 1826 

1826. Simia concolor Harlan, J. Acad. Nat. Sci. Philadelphia, 5, 4: 231, pis. g and 10. 

Locality unknown. (Hainan or Tonkin, Pocock, 1927.) 

1827. Hylobates harlani Lesson, Bull. Sci. Nat. Paris, /jj.- iii. Substitute for concolor. 
1840. Hylobates niger Ogilby, P.Z.S. 21. Error for concolor. 

1884. Hylobates nasutus Kunkel d'Herculais, Sci. et. Nat. 2: 86. Near Along Bay, 

Tonkin, Indo-China. 
1892. Hylobates hainanus Thomas, Ann. Mag. N.H. 9.- 145. Hainan. 
1897. Hylobates henrici Pousargues, Bull. Mus. H.N. Paris, i\- 367. Lai-chau, Tonkin. 
Range: Tonkin and Hainan. 


Hylobates concolor leucogenys Ogilby, 1840 

1840. Hylobates leucogenys Ogilby, P.Z.S. 20. Siam. Range: Siam and Laos. 

Hylobates concolor gabriellae Thomas, 1909 

1909. Hylobates gabriellae Thomas, Ann. Mag. N.H. 4: 1 12. Langbian, 1,500 ft., near 
Nha-trang, 100 km. inland from Phanrang, Southern Annam. 

Subgenus Sl'MPHALANGUS Gloger, 1841 

Hylobates syndactylus Raffles, 1821 Siamang 

Approximate distribution of species: Malay States, Sumatra; Tenasserim (accord- 
ing to Tate, 1947, Mamm. Eastern Asia). 

(Hylobates syndactylus syndactylus Raffles, 182 1. E.xtralimital) 

1821. Simia syndactyla Raffles, Trans. Linn. Soc. London, i^: 241. Bencoolen, 

Hylobates syndactylus continentis Thomas, 1908 

1908. Symphalangus syndactylus continentis Thomas, Ann. Mag. N.H. 2: 301. Semangko 
Pass, 3,000 ft., Selangor-Pahang border, Malay States. Range: northwards 
to Tenasserim ? 


For the continued use of Pholidota Weber, 1904, in spite of its preoccupation in 
the Reptilia, see Simpson (1945, 195). 

FAMILY: Manidae 

For a classification of this family see Pocock, 1924, The External Characters of the 
Pangolins, Manidae, P.Z-S- 707-723, with keys to all living subgenera. Pocock refers 
the seven existing species of Asia and Africa to six genera and three subfamilies. 
Simpson (1945) refers them all to a single genus. ^Vhile not denying the importance 
and interest of Pocock's work, Simpson's arrangement has much to commend it. 
Chasen appears to be in agreement, as he ignores Pocock's genus Paramanis. G. Allen 
follows Pocock. AVe do not consider Phatages valid even as a subgenus. 

Genus : Manis, page 2 1 4 



Genus MANIS Linnaeus, 1758 

1738. Manis Linnaeus, Syst. N'at. loth cd. /; 36. Manis pentadactyla Linnaeus. 
1762. PhoUdotus Brisson, Regn. Anim. 18. Based on Manis pentadactyla Linnaeus. 
1 81 5. Panaolinus Rafincsque, Analyse, 57. No type. 
■182 1. Pangolinii.s Rafincsque, Ann. Sci. Phys. Btux. 7; 214. Manis pcnladactvla 

1843. Phatages Sundcvall, K. Svenska Vetcnsk. Akad. Handl. 1842: 258, 273 (vel 

Phalagcnui). Mani': laticanda IlHger — Manis crassicaudata Gray. 
1873. Pangolin Gray, Handlist Edentate, etc., Mamni. Brit. Mus. 8. Based on Manis 

pentadactvla Linnaeus. 
1924. Paramanis Pocock, P.Z.S. 722. Manis javanica Desmarest. Valid as a subgenus. 

There are other, extralimital (African) subgencric names. 
3 species in Asia : 

Manis crassicaudata, page 215 
Manis javanica, page 215 
Manis pentadactyla, page 214 

A key to these species is given by Pocock (1924). 

Subgenus MANIS Linnaeus, 1758 

Manis pentadactyla Linnaeus, 1758 Chinese Pangolin 

Approximate distribution of species: Formosa, Southern China from Yunnan 
eastwards to Fukien, north to Kiangsu, and including Hainan; Burma, westwards to 
Sikkim and Nepal; Indo-China. 

Manis pentadactyla pentadactyla Linnaeus, 1758 

1758. Manis pentadactvla Linnaeus, Syst. Nat. loth cd. /.■ 36. Formosa. 

1777. Manis hrachrina Erxieben, Regn. An. ()8. 

^L\^TS PENTAD..\CTYL.\ .AURIT.A HodgSOn, 1 836. 

1836. Manis auntus Hodgson, J. As. Soc. Bengal, -,: 234. Lower and C^cntral Nepal. 

1843. Manis dalmanm Sundevall, K. Vet. Akad. Handl. Stockholm, 1842: 256, 278, 
pi. 4, fig. 10. Near Canton, Southern C^hina. 

1872. PhoUdotus assamensis Fitzinger, S.B. Akad. Wiss. Wien, 57. 

1872. Phatages hengalensis Fitzinger, loc. cit. 72. 

1907. PhoUdotus krevenbergt NLusthie, Wiss. Ergcbn. Exped. Filchner to C^hina, 10, i : 
234. Nanking, Kiangsu, China. 

Range includes Nepal, Sikkim, Naga Hills in Assain (B.M.), Pegu and Mt. Poppa in 
Burma, Laos, Tonkin, and Yunnan to Fukien, Anhwci, Kiangsu, etc., in Southern 
China. G. Allen called this race M. p. dalmanui, with aurila in the synonymy, but 
aurita takes priority by seven years. 


1 006. Manis pusilla ]. Allen, Bull. Amcr. Mus. N.H. 22: 46-,, pi. 69, figs. 1^3. Island 
ol Hainan. 



Manis crassicaudata Gray, 1827 Indian Pangolin 

Approximate distribution of species: Ceylon, Peninsula of India (Shevaroy Hills, 
Madras, Mysore, Bellar^', Kanara, Coorg), to Cutch and Bengal. (Blanford (i8gi) 
who erronoeusly called this species M. pentadactyla, said it occurred in Peshawar, 
Sind and Orissa.) G. Allen thought its range extended to extreme Western Yunnan. 

Manis crassicaudata Gray, 1827 

1815. Manis laticauda Illiger, Abhandl. Preuss. Akad. VViss. 1 804-1 811; 90, nom. nud. 

1827. Manis crassicaudatus Gray, in Griffith's Cuvier Anim. Kingd. 5; 282. India. 

It is customary to date the name crassicaudata from Geoffroy, 1803, Cat. Mamm. Mus. 
H.N. Paris, 213, but according to Sherborn this work was never published. 

1865. Pholidotus indicus Gray, P.Z.S. 368. 

Subgenus PARAMANIS Pocock, 1924 

Manis javanica Desmarest, 1822 Malayan Pangolin 

Approximate distribution of species: Burma, Tenasserim, Indo-China, Siam, 
Malay States, Sumatra, Java, Borneo, many small adjacent islands, east to the 

Manis javanica Desmarest, 1822 

1822. Manis javanica Desmarest, Ency. Meth. Mamm. 2: 377. Java. 

1842. Manis leplura Blyth, J. Asiat. Soc. Bengal, //.• 454. Locality unknown. 

1847. Manis leucura Blyth, J. Asiat. Soc. Bengal, 16: 121^. Arakan, Burma. 

1850. Manis guy Focillon, Rev. Mag. Zool. 2: 513, pi. 10. Locality unknown 

Range: as above, in Indo-China, including Laos, Annam, Cochin-China. 


Among special works of reference to this Order are: 

Miller, G. S. 1912. Catalogue of the Mammals of Western Europe. 

Allen, G. M. 1938. Mammals of China & Mongolia, Natural History of Central .Asia, 11: 
I. New York (American Museum of Natural History). 

1939- A Checklist of African Mammals. Bull. Mus. Comp. ^ool. Harvard, 8j. 

Pocock, R. I. 1939, 1941. The Fauna of British India, Mammals, i and 2; and numerous 
short papers. 

Bobrinskii, N., Kuznetzov, B. & Kuzyakin, A. 1944. Mammals of the U.S.S.R. 

Simpson, G. G. 1945. The Principles of Classification and a Classification of Mam- 
mals. Bull. Amer. Mus. N.H. 8§. 

Ognev, S. I. 1931, 1935. Mammals of Eastern Europe and Northern .isia, 2 and j. 


Simpson 1045) cli\idcs li\ing members nf this Order into two superfamilies: 
Canoidea ^ containing the families Canidac, Ursidae, Procyonidae and Mustclidae) ; 
and Feloidca (containing the families Vivcrridae, Fehdae and Hyaenidae). These 
superfamilies correspond to the suborders Aeluroidea and Arctoidea of Pocock 
(1041), and other authors. \Ve prefer to follow Simpson and regard these two groups 
as of superfamily rank. The classification of Simpson is simpler than that of Pocock, 
and more conservativa. It is here followed, with some small generic modifications. 

Neither Simpson nor Pocock give the Seals (Pinnipedia) ordinal rank. Simpson 
(p. 121) lists them as a suborder, and Pocock considered them as part of the 
"Arctoidea". However, other authors, as Miller, G. Allen, Ognev and Bobrinskii 
treated the Pinnipedia as a distinct order. Simpson (p. 232) seems to suggest that the 
group is an old one, widely separated from the Carnivora as here understood, and 
the convenience of giving the group ordinal rank seems so marked that we here 
follow Miller and others, and regard the Pinnipedia as an order distinct from the 

FAMILIES: Canidac, page 216 
Felidae, page 300 
Hyaenidae, page 299 
Mustelidae, page 243 
Procyonidae, page 242 
Ursidae, page 235 
Viverridae, page 279 


Genera: Alopcx, page 222 

Canis, page 2 1 7 

Ciwn, page 233 

Fennecus, page 231 

Lycaon, page 234 

Nvctereutes, page 222 

Vulpes, page 223 
Simpson dix'ides existing Canidae into three subfamilies, one of which, the 
Otocyoninac, is extralimital and doubtless valid. The Cuoninae, or Simocyoninae as 
listed by Simpson, containing Cuon and Lycaon, is not supported by Pocock, 1941, 
2: 146. 

We know of no paper which specially compares the various genera of Canidae 
with each other. Our translation of Ognev's key to the genera of Canidae in the 
U.S.S.R. indicates that in Nxcterades the posterior edge of the mandible has a lobate 
process separated by a notch from the markedly elevated angular process, the latter 
being short, round, and indistinctly separated from the condylar process by a shallow 
hollow, thereby differing from the mandibles of Canis, Vulpes and Alopex (and (in 
B.M. material) also from that of Fninceus). Bobrinskii (p. 139) gives a figure of the 
skull of ,,\V(7(7V7//(y, which may be compared with Miller's figures o( Canis, Vulpes and 



Alopex. There are also external differences, such as the short ears, and rather short 
limbs, by which Nyctereutes may be separated from Canis, etc. Generic characters of 
Canis, Vulpes and Alopex are given by Miller (191 2, 304); and those of Canis and 
Vulpes are compared with Cuon by Pocock (1941, 80). Fennecus is like a small Vulpes, 
but with enormous bullae and ears. Pocock did not retain it as a genus, but there 
seems httle doubt that it should be retained. It antedates Vulpes. Lycaon is largely 
extralimital, but is included on the basis of a note in G. Allen (1939) on' skulls from 
Tanezrouft, Algeria, which is within the North African Pala'earctic ; it differs from 
the other Palaearctic genera in the suppression of the pollex, and is very different 
from the others in general appearance, its characters including spotted body, large 
rounded ear, and relatively very large size. 

Mivart, i8go. Monograph of the Canidae, still seems to be the best general work on 
this family. There are good figures of all the leading species, but it is out of date in 
some ways, for instance as regards genera now recognized. 

Genus CANIS Linnaeus, 1 758 

1758. Canis Linnaeus, Syst. Nat. loth ed. /.• 38. Canis familiaris Linnaeus (the 

domestic dog). 
1816. Thos Oken, Lehrb. d. Naturgesch. j, 2: 1037. Thos vulgaris Oken = Canis 

aureus Linnaeus. 
1816. Lupus Oken, Lehrb. d. Naturgesch. 3, 2: 1039. Canis lupus Linnaeus. 
1837. Vulpicanis Blainville, Ann. Sci. Nat. Paris, Zool. 8, 2 : 279. Canis aureus Linnaeus. 
1839. Sacalius H. Smith, Jardine's Naturalists Library, Mamm. 2j: 214. Sacalius 

aureus [Canis aureus Linnaeus). 
1841. Oxygous Hodgson, Calcutta, J.N. H. 2: 213. Canis aureus Linnaeus. 
1855. Lupulus Gervais, H.N. Mamm. 2: 60-62. Not Lupulus Blainville, 1843. 
1869. Dieba Gray, Cat. Carn. Pachyd. & Edentate Mamm. B.M. 180. Canis anthus 

F. Cuvier. 
1906. Lupulella Hilzheimer, Zool. Beobachter, 4j: 363. Canis mesomelas Schreber. 
1906. Schaeffia Hilzheimer, Zool. Beobachter, ^7.- 364. Canis adustus Sundevall. 
1906. Alopedon Hilzheimer, Zool. Beobachter, 4y: 365. Canis thooides = Canis anthus 

Cretzschmar nee Cuvier. 

2 species in the area covered by this list : 
Canis aureus, page 220 
Canis lupus, page 2 1 8 

For the characters of the two Palaearctic species see Miller (19 12, 305) and 
Pocock (194 1, 82). For a note on the characters of the three widely-distributed 
African species of Jackals, C. aureus, C. adustus Sundevall, 1846, and C. mesomelas 
Schreber, 1778, see Hollister, 1918, Bull. U.S. Nat. Mus. gg: loi. Hilzheimer, 
in 1906, named a subspecies of Jackal Canis lupaster grayi, from Morocco and Tunis, 
and G. Allen, in his Checklist of African Mammals, for no apparent reason, lists this 
form as a race of the otherwise Ethiopian species Canis adustus. Hilzheimer said 
that his race was the same as that figured by Gray, 1868, P.^-S. 503. This figure 
is oi Canis aureus subsp. It bears no close resemblance to the skull oi Canis adustus, and 
there is little evidence that adustus occurs in any part of Palaearctic North Africa. 


p.\i.aearc;tic and Indian mammals 1758-1946 

Canis lupus Linnaeus, 1758 \Volf 

Approximate distribution of species: formerly widely distributed in Europe, in- 
(.iudins; the British Isles, but now extinct in Western Europe except for Portugal, 
Spain, Italy, Sicily, Sweden and (occasionally) Norway. Widely distributed in the 
U.S.S.R. The western limit of the Russian wolves fluctuates considerably, since the 
animals are much given to wandering, but may be taken as a line running from 
Sweden, through Finland, and then along the eastern borders of the Baltic States, 
East Prussia, Poland and Czechoslovakia; thence through Rumania to Yugoslavia 
and Bulgaria, with occasional extensions into Northern Greece and Turkey. The 
Asiatic range includes,' according to Bobrinskii, Russian Asia ("all over the Union, 
except Crimea and \arious northern islands, but inhabits Sakhalin, Bolshoi Lyakhov- 
skii Island, the south island of Novaya Zemlya and Kolgucv") ; Mongolia, Korea, 
Japan (if not extinct there), Tibet; Kansu, eastwards to Chihli in China (perhaps, 
also other parts of China) ; in India, from Baluchistan and Kashmir southwards, at 
least to Dharwar, and eastwards to Bengal, and in South-A\'cstern Asia, from Persia, 
Iraq, Asia Minor, Palestine and Arabia. Widely distributed in North America. 
For review, sec Pocock, 1935 P-Z-S- 647. 

Cams lupus lupus Linnaeus, 1758 

1758. Canis lupus Linnaeus, Syst. Nat. loth cd. /; 39. Sweden. 

1792. Cams lupus flavus Kerr, Anim. Kingd. 137. France and Germany. 

1804. Canis lupus niger Hermann, Obscrv. Zool. 32, not of Kerr, 1792. Forest of 

Hagcnau, Alsace. 
1804. Cajiis lupus communis Dwigubski, Pn>d. Faun. Ri;ss. 10. Russia. 
1839. Cams lupus var. canus dc Selys Longchamps, Etudes de Micromamm. 144, 

nom. nud. 
1839. Cams lupus var. Julvus dc Selys Longchamps, luc. cil., nom. nud. 
1841. Lupus onentalis Wagner, Schreb. Saugeth. Suppl. 2: 3G7. Europe. 
1863. Canis lupus var. major Ogerien, H.N. du Jura, j: 64. Lower slopes of the Jura. 
1863. Canis lupus var. minor Ogerien, loc. cil. Higher portions of the Jura. 
1910. Canis lupus lycaon Trouessart, Faune Mamm. Europe, 90. Pyrenees. 
(?) 1911. Lupus aliaicus Noack, Zool. Anz. 35: 465. Chulyshman Glacier, Altai. 
(?) 1922. Canis lupus var. onentalis Dybowski, Arch. Tow. Nauk. Lwow, /.• 350, nom. 

nud. Xec \Vagner, 1841. 
!?) 1922. (auiis lupus var. argunensis Dybowski, loc. cil., nom. nud. 
Range: Northern and Clentr.i! Europe, and forest zone of the Li.S.S.R. 

Canis lupus albus Kerr, 1 792 

1792. Canis lupus albus Kerr, Anim. Kingd. 137. Near Jenisca, in the eastern part of 

Asiatic Russia. 
(.^J i<)22. Canis lupus var. kamlschalicus Dybowski, .\rch. Tow. Nauk. Lwow, /.• 350. 

Kamtchatka, nom. nud. 
ir)23. Ganis (sic) lupus luruchanensis Ognev, Biol. Mitt. TimiriazelF, /; 113. Turuk- 

hansk region 'on Northern Venesei), Siberia. 
1?) 1926. Cams lupus dvbowskii Domanicwski, Pracc Z. Mus. Warsawa, 5.- 52. Goly- 

gina, South-\\'estern Kamtchatka. 
Range: whole tundra and liirest-tundra area of LLS.S.R. 



Canis lupus campestris Dwigubski, 1804 

1804. Canis lupus campestris Dwigubski, Prod. Faun. Ross. 10. In deserts between 

Black Sea and Caspian, Kirghizia, to River Yenesei. 
(?) 1882. Canis lupus var. desertorum Bogdanov, \.H. Khibinsk Oasis and Desert 

Kizilkum, 30. {N.V.) Kizil Kum Desert, Russian Turkestan. 
(?) 1923. Canis lupus cubanensis Ognev, Biol. Mitt. Timiriazeff, /; 1 14. Maikop district, 

Kuban region, Southern Russia (Caucasus). 

Bobrinskii lists only one subspecies of C. lupus from the deserts and steppes of 
Central Asia and Kazakstan, which he calls C. /. desertorum, but it would seem that 
campestris Dwigubski antedates. 

Canis lupus pallipes Sykes, 1831 

1 83 1. Canis pallipes Sykes, P.Z.S. loi. Deccan, India. Range: the plains of Northern 

India from Bengal to Sind, south to Dharwar, also Baluchistan, and thence 

westwards to Iraq and Northern Arabia. 

Canis lupus hodophilax Temminck, 1839 

1839. Canis hodophilax Temminck, Tijdschr. Natuurl. Geschied. Physiol. 5.- 284 (see 

Harper, 1940. J. Mammal. 2j: 192). Hondo, Japan. 
1844. Canis hodopylax (sic) Temminck, Fauna Japon. Mamm. 38, pi. 9. Nippon or 

Hondo, Japan. 
1885. Canis lupus japonicus Nehring, S.B. Ges. Nat. Fr. BerHn, 141. 
Range: Hondo, Japan (said to be extinct, Kuroda, in Harper (1945) ). 

Canis lupus chanco Gray, 1863 

1863. Canis chanco Gray, P.Z.S. 94. Chinese Tartary. 

1847. Lupus laniger Hodgson, Calcutta J.N. H. j: 474. Tibet. Not C. laniger H. Smith, 

1874. Canis niger Sclater, P.Z.S. 655, pi. 78. Not of Kerr, 1792. Hanle, Kashmir. 
1883. Canis ekloni Przewalski, Third Journey to Tibet, 216, nom. nud. 
1907. Lupus filchneri Matschie, in Filchners Exped. to China, \Viss. Ergebn. 10, i: 

153. Siningfu, Kansu, China. 
1907. Lupus karanorensis Matschie, loc. cit.: 159. Karanor, in the Gobi. 
1907. Lupus tschiliensis Matschie, loc. cit.: 160. Coast of Chihli, China. 
1923. Canis lupus coreanus Abe, Dobuts. Zasshi. 55.- 383. Onpeimen, near Seoul, in 

Keikido Province, Korea. 

Range: Russian Pamir, Chinese Turkestan, Tianshan, Tibet, Mongolia, Northern 
China (including Shensi). 

Canis lupus signatus Cabrera, 1907 

1907. Canis lupus signatus Cabrera, Bol. Real. Soc. Esp. H.N. Madrid, j: 195. Escoril, 
Madrid, Spain. 

Canis lupus deitanus Cabrera, 1907 

1907. Canis lupus deitanus Cabrera, Bol. Real. Soc. Esp. H.N. Madrid, 7: 197. Mora- 
talla, Murcia, Spain. 

P 219 

PALAEARC:TIC: and IXDIAX mammals 1758-1946 

CIanis Lupfs iTALicus AltobcUo, 1921 

iq2i. Canii lupus italicus Altobcllo, Fauna DcU'Abruzzo del Molisc, Mammiferi, ^; 
41. Abruzzi, Italy. 

CIanis lupus kurjar Bolkay, 1925 

1925. Cams lupus kurjak Bolkay, Nov. Mus. Saraje\'o, Xo. i, 9. Teslie, Bosnia, 

Cams lupus h.'^ttai Kishida, 1931 

1931. Canis lupus hatlai Kishida, Lansania, jj, 25: 73. 'A'.I'.) City of Sapporo, 

Hokkaido, Japan. 
1935. Canis lupus rex Pocock, P.Z.S. 659. Yezo (= Hokkaido). Extinct in Hokkaido, 

but sun-ivint; in .Sakhalin and perhaps in the Kuriles (Harper, 1945). 

Canis lupus ar.^bs Pocock, 1934 

1934. Canis lupus arahs Pocock, Ann. .Mag. N.H. i ^: 636. Ain, Southern Arabia, 
1,500 ft. 

Canis aureus Linnaeus, 1758 Asiatic Jackal 

Approximate distribution of species: Balkan States, Rumania, Greece; Russian 
Turkestan (Western and Southern Turkmenia, Tadzhikistan, whole course of Amu- 
Darya, Samarkand and Bokhara districts. Middle Syr-Darya), Persia, Iraq, Asia 
Minor, Afghanistan 'according to Bobrinskii), Syria, Palestine, Arabia; Baluchi- 
stan and Sind, south through Peninsular India to Ceylon, eastwards to Nepal, 
Assam, Burma and Siam. Egypt, Libya, westwards to Morocco, Rio de Oro, thence 
southwards to Senegal, the .Sudan, Somaliland, Abyssinia and Kenya. 

C.^Nis aureus aureus Linnaeus, 1758 

1758. Cams aureus Linnaeus, Syst. Nat. lOth cd. /.■ 40. Proxince of Lar, Persia. 

1 84 1. C anis) aureus vulgaris Wagner, Schreb. Saugeth. Suppl. i^.- 383. 

(?) 1841. Canis dalmatinus Wagner, Schreb. Saugeth. Suppl. -'.■ 383. Dalmatia. 

1858. Canis aureus typicus or var. caucasica Kolenati, Reiseerinerungen, /.' 96. 

(?) 1892. Canis aureus balcanicusBrwim^i, Glasnik Hrvatskoga Naravoslovnoga Drustva, 
Zagreb, /.■ 317. Drava River, C^roatia. See Pocock, 1938, P.Z.S. , Ser. B. 108: 
37, 39, in which it is sugge^tcd that dalmatinus and balcanicus are possibly 
synonyms of C. a. anthus Cuvier, 1820, from Senegal, evidently introduced 
into Europe. 

1896. Cams hadramauticus Noack, Zool. .-\nz. if/: 356. .\rabia. Noack's species is a 
composite one made from a jackal and a wolf; the jackal was chosen as 
Icctotype by Matschie (sec Xlorrison-Scott, 1939, Mov. Zool. 41: 201). 

1916. Canis indicus kola Wroughton, J. Bomljay N.H. Soc. 2^: 651. Palanpur, 
Gujerat, Western India. 

Range: Iraq, Persia, Baluchistan, Western India Clutch, Sind, Gujtrat), Arabia, 


Canis aureus syriacus Hemprich & Ehrenberg, 1833 

1833. Canis syriacus Hemprich & Ehrenberg, Symb. Phys. Mamm. text 2, sig. z, 

pi. 16. Coast of Lebanon, between Beirut and Tripoli. Range: Syria, 


Canis aureus lupaster Hemprich &. Ehrenberg, 1833 

1833. Canis lupaster Hemprich cS: Ehrenberg, Symb. Phys. Mamm. 2, sig. ff. Fayum, 

1833. Canis sacer Hemprich & Ehrenberg, Symp. Phys. Mamm. 2, sig. ff. Fayum, 

Range: Egypt, Palestine (part), according to Bodenheimer, and Libya. 

Canis aureus indicus Hodgson, 1833 

1833. Canis aureus indicus Hodgson, Asiat. Res. 18, 2: 237. Nepal. Range: Nepal, 
Sikkim, Bhutan, Assam, Burma, Siam. 

Canis aureus moreoticus L Geoffroy, 1835 

1835. Canis aureus var. moreotica Geoffroy, Exped. Sci. de Moree, Zool. pi. i. Morea, 

1 84 1. Canis ^raecus Wagner, Schreb. Saugeth. Suppl. 2: 383. Peloponesus, Greece. 
Range: Greece, Asia Minor and Caucasus (Pocock, who used this name for the 
European jackals). 

Canis aureus algirensis Wagner, 1841 

1839. Sacalius barharus H. Smith, Nat. Lib. Jardine Mamm. 25.- 218. Tunis. Not of 

Shaw, 1800. 
1841. Canis aureus algirensis Wagner, Schreb. Saugeth. Suppl. 2: 384. Algeria. 
1841. Canis aureus tripolitanus \V'agner, loc. cit. No locality; Tripoli, Tunis implied. 
(?) 1906. Canis lupaster grayi Hilzheimer, Zool. Beobachter, ^j: 367. Morocco and 

1906. Canis studeri Hilzheimer, Zool. Beobachter, 4J: 368. Tunis. 

Canis (?) aureus cruesemanni Matschie, 1900 

1900. Canis cruesemanni Matschie, S.B. Ges. Nat. Fr. Berlin, 145. Menam, Siam. 
Status doubtful; based on living captive specimens. 

Canis aureus soudanicus Thomas, 1903 

1903. Canis aureus soudanicus Thomas, P.Z.S. /.• 295. El Obeid, Kordofan, .Sudan. 
(?) 1826. Canis variegatus Cretzschmar, in Riipp. Atlas Reisc nOrdl. Afrika, Saugeth. 31, 

pi. 10. Not Canis familiaris variegatus Gmelin, 1788. Nubia and Upper Egypt. 
1906. Canis doederleini Hilzheimer, Zool. Anz. 30: 1 16. Upper Egypt. 
1921. Thos aureus nubianus Cabrera, Bol. Real. Soc. Esp. H.N. Madrid, 21: 264. To- 

replace variegatus Cretzschmar, preoccupied. 

Canis aureus naria Wroughton, 19 16 

1916. Canis naria Wroughton, J. Bombay N.H. Soc. 24: 651. Virajpet, Southern 
Coorg, 3,000 ft., India. Range: Southern Peninsular India. 


Canis aureus LANKA Wroughton, 1916 

1916. Canis lanka Wroughton, J. Bombay N.H. Soc. 24: 652. Mankeni, East Pro- 
vince, Ceylon. 

Canis aureus maroccanus Cabrera, 1921 

1921. Thos lupaster maroccanus Cabrera, Bol. Real. Soc. Esp. H.N. Madrid, :'/.• 263. 
Mogador, Morocco. 

C.-^Nis aureus ecsedensis Kretzoi, 1947 

1947. Thos aureus ecsedensis Kretzoi, Ann. Mus. Nat. Hung, ^o: 287. Tyukod, 

Szatniar, Hungary. Proposed to replace huns,aricus. 
1938. Canis aureus hungaricus Ehik, Ann. Mus. Nat. Hung, jj/ (Zool.) : i i. Said to be 

preoccupied by Canis fanuliaris hungaricus Margo, 1891, the reference to 

which has not been traced. 
1897. Canis lupus minor Mojsisovico, Das Thicrleben d. Ost. Ung. Tielebenen, 244. 

Northern Hungary. Said to be preoccupied by Canis spelaeus minor Wagner, 

1 83 1, the reference to which has not been traced. Not of Ogerien, 1863. 

Genus ALOPEX Kaup, 1829 

1829. Alopex Kaup, Skizz. Europ. Thierw. /; 83, 85. Canis lagopus Linnaeus. 
1868. Leucoevon Gray, P.Z.S. 521. Canii lagopus Linnaeus. 

Bobrinskii, 1944, Manimali U.S.S.R. 146, regards Alopex as a subgenus of Vulpes. 

I species: Alopex lagopus, page 222 

Alopex lagopus Linnaeus, 1 758 Arctic Fox 

Approximate distribution of species: Norway, Sweden, Spitzbcrgcn, Iceland, 
Arctic regions of LI.S.S.R., from European Rtissia to Kamtchatka and the Pacific, 
and perhaps south to Kurile Islands; also in Northern North America. 

Alopex l.xgopus lagopus Linnaeus, 1 758 

1758. Canis lagopus Linnaeus, Syst. Nat. loth ed. /.• 40. Lapland. 

1816. Vulpes arclica Oken, Lehrb. d. Naturgesch. j, 2: 1033. 

1820. Canii vulpes caerulea Nilsson, Skand. Fauna, /.■ 88. Lapland. 

1827. [Canis lagopus) argcnteus Billberg, Synop. Faunae Scandinaviae, 14. Lapland. 

1898. Canis lagopus typicus Barrett-Hamilton & Bonhote, Ann. Mag. N.H. /; 287. 

Range: apparently the mainland range of the species. 

.-Xlijpex lagopus fuliginosus Bechstein, 1799 

1709. Canis fuliginosus Bechstein, Thomas Pennants allgem. Uebersicht d. vierf. 
Thierc, /.' 270. Iceland. A\ailablc if the Iceland race proxies distinguishable. 

Alopex laoopus spiTZBERCiENENsis Barrett-Hamilt(in & Bonhote, 1898 
1898. C.ani^ lagopus spilzhergenensis Barrctt-Hamiitnn & Bonhote, Ann. Mag. N.H. i: 
2K7. Spitzbcrgcn. 


Alopex lagopus beringensis Merriam, igo2 

igo2. Vulpes beringensis Merriam, Proc. Biol. Soc. Washington, /j; 171. Bering Island, 

Bering Sea, Eastern Siberia. 
1920. Alopex beringianus Cherski, Komandorskinesez, Tokyo, 60 [M.V.) 

Genus VULPES Oken, 1816 

1775. Vulpes Frisch, Natur-system der vierfuss. Thiere, 15 (see page 3). 

1816. Vulpes Oken, Leiirb. d. Naturgesch, j, 2: 1033, 1034. Vulpes vulgaris Oken = 

Canis vulpes Linnaeus (see page 3). 
1822. Vulpes Fleming, Philosophy of Zool. Edinburgh, 2: 184. Canis vulpes Linnaeus. 
1839. Cynalopex H. Smith, Jardine's Nat. Library, Mamm. 25.- 222. Canis corsac 


6 species in the area covered by this list: 

Vulpes bengalensis, page 230 Vulpes ferrilala, page 231 
Vulpes cana, page 231 Vulpes riippelli, page 230 

Vulpes corsac, page 229 Vulpes vulpes, page 225 

In an attempt to correlate the work of Pocock, 1941, Fauna Brit. India, 2: 1 10; 
Miller, 191 2, Cat. Mamm. IV. Europe; Bobrinskii, 1944, Mamm. U.S.S.R.; and G. 
Allen, 1938, Mamm. China & Mongolia, and to add notes on the outlying forms of 
the genus from Africa, South-Western Asia and Japan, the following results have 
been obtained: 

1. Back of the ears black or dark brown, contrasting strongly with colour of head 

and nape. VULPES VULPES 

(Forms available for examination: karagan, crucigera, aegyptiaca, montana, 
atlantica, flavescens, pusilla, griffithi, japonica, hoole, beringiana, arabica, silacea, 
induta, ichnusae, anatolica, palaestina.) 
Back of the ears generally same colour as the head and neck, never strongly 

contrasted. 2 

2. Tail less than half head and body length; ear less than or equal to half the length 

of the hindfoot (according to the published measurements of Pocock, G. Allen 

and Mivart). 3 

Tail clearly more than half length of head and body (normally). Ear clearly more 
than half length of hindfoot. 4 

3. Skull much larger; bullae appear larger; muzzle long and narrow; upper canine 

elongated, clearly larger than combined length of P 4 and M i in upper jaw. 

Skull considerably smaller; bullae appear smaller; muzzle not specially elongated 
nor narrow; upper canine scarcely or only a little exceeding combined length of 
P 4 and M I in upper jaw. VULPES CORSAC 

(Not well represented in London: three skulls only and a few unmeasured 



4. Tail tip clearly contrasted white; cr, in the case (one specimen availablcl of" 
Ziiiiidim. \sh('iie tail appears whitish. WIPES RUPFELU 

(Forms a\ailable for examination: >ii/i/>flli, caesia, zo'i'dim, iabaen, somaliae 
(Thomas, 11)18, from Somaliland).) 
Tail tip normally clearly contrasted black; never sharply contrasted white. 


5. Larger species; head and body length, with few exceptions, not less than 

460 mm. 'j 

Smaller species; head and body length in the majority of specimens does not 
exceed 420 nmi. 7 

6. Ear length normally 85 mm. and more. VULPES CHAMA (Smith, 1833) 

(Extralimital; from South Africa.) 
Ear length 84 mm. and less, but in the very considerable series in the British 
Museum, only three specimens as long as 81 mm. VULPES BENGALENSIS 

7. Fur very thi( k; darker in colour; a dark middorsal line traceable in all .skins; 

black tailtip weaker. Ear (of one skin) 88 mm. VULPES CANA 

Fur thin and short; colour pale; no middorsal line; black t.iiltip normally \-cry 
sharply contrasted. Ear not exceeding 75 mm. in British Museum skins. 
(Extralimital) VULPES PALLIDA Clretzschmar, 1826 

(Forms a\ailable lor examination: pallida, Sudan; edwardsi, Rochebrune, 
1833, Senegambia; and /((7/7<77; Thomas & Hinton, 1921, Northern Nigeria.) 

Measurements in the above key for V. cana and V. ferrilata are mainly based on 
those given by Pocock (1941). There is very little data on exact measurements of 
['(///'(■) (-01 sac. which is the second name in the genus. Measurements given by G. 
Allen, and Misart, suggest that it is correctly placed in the above key. In appear- 
ance, hf!ii;al(')i^!\ is not very widely separated from it. It is, according to Boljrinskii, a 
larger animal than V. cana. This author notes it as with ears and tail comparatively 
short. Viil/:f^ liiiiliila seems in some ways the most distinct of the species. Its dental 
and cranial characters given in the key contrast with all other Indian species. I', vulpes 
is at extreme development the largest species. V. li/ppelli has large ears, 80 mm. at 
lowest, and up to wjo mm. in British Museum material. Normally it is larger than 
pallida, but the Arabian race may sometimes be an exception. It occurs in the same 
general neighbourhood ?i% pallida, and compared with its immediate allies its white 
tailtip seems \ery distinctive. We can trace no fox in Central Tropical .Africa; that is 
to say, south of the Senegal-Northern Nigeria-Sudan-Somaliland line; north of 
Angola and South-\\cst Ai'rica. (The British Museum possesses I'ulpa skins Inim 
Angola.) ['. cfiama seems geographically isolated in 'the south. In Africa, V. vuljja is 
strictly Palaearctic. The form dorsalis listed by G. Allen from Senegal is a jackal, 
pr<ibabh' Caiiis aureus; ly|)<' skin in British Museum. 

G. Allen, H| ;i), listed the Libyan form cvrenaica as a race of I', pallida, but from the 
description it is much more likely that it represents V riippelli. 



Vulpes vulpes group 

Vulpes vulpes Linnaeus, 1758 Common Red Fox 

Approximate distribution of species: essentially throughout the Palaearctic region; 
in South-Eastern Asia south of it into Yunnan, Fukien, and Northern Indo-China; 
and, according to Pocock, also much of North America. 

(In detail: British Isles, Ireland included; France, Belgium, Holland, Denmark, 
Norway, Sweden, Germany, Switzerland, Spain, Portugal, Italy, Sardinia, Poland, 
Rumania, Greece, doubtless other European countries; the whole of the U.S.S.R. 
("but it apparently does not penetrate into the interior of the tundra, and fails to 
occur in the extreme north of Siberia and on nearly all the islands of the Arctic 
Ocean and Bering Sea, only appearing on Kolguev Island and the south island of 
Novaya Zemlya; occurs in Sakhalin" (Bobrinskii) ) ; Arabia, Persia, Afghanistan, 
Cyprus, Palestine, Iraq, Asia Minor; Western Sinkiang (Ognev), Mongolia, Japan, 
Manchuria, Tibet, and the states of Yunnan and Fukien northwards in China; 
India, from Rajputana, Sind, Cutch and Khandesh, northwards to Baluchistan, 
VVaziristan, Punjab, Kashmir, Sikkim; Tonkin, in Indo-China; Egypt, Algeria, 
Libya and Morocco.) 

Vulpes vulpes vulpes Linnaeus, 1758 

1758. Canis vulpes Linnaeus, Syst. Nat. loth ed. /.■ 40. Sweden. 

1758. Canis alopex Linnaeus, Syst. Nat. loth ed. /.• 40. Sweden. 

1816. Vulpes vulgaris Oken, Lehrb. Nat. 3, 2: 1034. 

1820. Canis nigro-argenteus Nilsson, Skand. Fauna, /.• 91. Lofoten Islands, Norway. 

1827. Canis vulpus nigrocaudaius Billberg, Synop. Faunae Scandinaviae, 12. Uppland, 

1827. Canis vulpus variegatus Billberg, loc. cit. 13. Uppland, Sweden. 
1827. Canis vulpus lineatus Billberg, loc. cit. 13. Skane, Sweden. 
1830. Vulpes communis Burnett, Quart. J. Sci. Lit. Art. iSsg, 2: 349, nom. nud. 
Range: Scandinavia. 

Vulpes vulpes karagan Erxleben, 1777 

1777. Canis karagan Erxleben, Syst. Regn. Anim. Mammalia, 566. Kirghiz Steppes, 

Russian Asia. 
181 1. Canis mclanotus Pallas, Zoogr. Ross. Asiat. /.• 44. 
1 926. Vulpes vulpes karagan nutio ferganensis Ognev, Ann. Mus. Budapest, 2j: 222. Osh, 

Fergana, Russian Turkestan. 
1926. Vulpes vulpes karagan natio pamirensis Ognev, loc. cit. Pamir Mountains. 
Range: Kirghiz and Kazakstan steppes, to Mongolia. 

Vulpes vulpes crucigera Bechstein, 1789 

1789. Canis crucigera Bechstein, Gemeinn. Nat. Deutschlands, /.• 250. Thuringia, 

1792. Canis vulpes alopex europaeus Kerr, Anim. Kingd. 142. Burgundy, France. 
1797. Canis vulpes alba Borkhausen, Deutsche Fauna, /.• 33. Vogelsberg, Hesse, 

Germany. Not of Kerr, 1792. 
1 797. Cams vulpes nigra Borkhausen, loc. cit. Hesse and Thuringia, Germany. 




1801. Canis vulpes lutea Bechstcin, Gemcinn. Nat. Deutschlands, /, 2nd ed. : 628. 

Thuringia, Germany. 
1801. Canis vulpes cincra Bechstein, he. eit. Thuringia, Germany. 

1832. Canis melanogaster Bonaparte, Iconogr. Fauna Ital. /.■ fasc. i . Near Rome, Italy. 
1 84 1 . Vulpes hrpomelas Wagner, Schreb. Saugeth. Suppl. 2: 405. Oberbayern, Germany. 
(?) 1855. Vulpes vulgaris meridionalis Fitzinger, Wissensch. pop. Nat. der Saugeth. i: 

U)4. Dalmatia. (Published in 1860 according to Miric, D., Z. Saugetierk. 
i960, 2-,: 45). 
Range: British Isles, France, Germany, .Switzerland, Italy, Sardinia, Greece, forested 
parts of Northern and Gentral Russia. 

Vulpes vulpes barb.'^ra Shaw, 1800 

1800. Canis harbarus Shaw, Gen. Zool. /, Mamm. pt. 2, 311. Barbary, i.e. coast of 

North-Western Africa. 
1916. Vulpes vulpes acaab Cabrera, Bol. Real. Soc. Esp. H.N. .Madrid, 16: 384. 

Marraquex, Western Morocco. 

Vulpes vulpes aegyptl\ca Sonnini, 1816 

1 8 16. Canis aegyptiacus Sonnini, Nouv. Diet. Sci. Nat. 6: 524. Egypt. 

1820. Canis niloticus Desmarest, Encyclop. Method. Mamm. 204. E.gvpt. 

1833. Cams anubis Hemprich & Ehrenberg, Symp. Phys. Mamm. dec. 2, sig. ff. 

Fayum, Egypt. 
1833. Canis vulpeeula Hemprich & Ehrenberg, loc. eit. Fayum, Egypt. 
Range: Egypt, Libya and Palestine (according to Bodenheimer). 

Vulpes vulpes Montana Pearson, 1836 

1836. Canis vulpes montana Pearson, J. Asiat. Soc. Bengal, 5; 313. (January, 1836.) 


1837. Canis himalaieus Ogilby, P.Z.S. 1836, 103. (20 February 1837.) Mussooree, 

Kumion, North-Western India. 
1837. Vulpes nepalensis Gray, Charlesw. Mag. N.H. /.• 578. Nepal. 
1888. Vulpes alopex Blanford, Mamm. British India, 153. Not of Linnaeus, 1758. 

1906. Vulpes waeidelli Bonhote, Abstr. P.Z.S. 14; P.Z.S. 303. Kambajong, Tibet. 

1907. Vulpes ladaeensis Matschie, AN'iss Ergebn. Filchners Exped. China, 10, i: 167. 

Range: Sikkim, Yunnan, Tibet, Kumaon, Nepal, Punjab, to Gilgit. 

Vulpes vulpes atlantica Wagner, 1841 

1841. Cams vulpes var. atlantica Wagner, Reisen in d. Regenschai't Algier, 5.- 31, 62, 

pi. 3. Atlas Mounta,ins, Mitiya, Algeria. 
i8-j8. Vulpes algeriensis Loche, Cat. Mamm. et Oiseaux observes en Algerie, 4. 

Wooded parts of Algeria. 

Vulpes vulpes flavescens Gray, 1843 

1843. Vulpes flavescens Gray, Ann. Mag. N.H. //.• 118. Northern Persia. 

1902. Vulpes vulpes splendens Thomas, Ann. Mag. N.H. 10: 489. Astrabad, Persia. 

1912. Vulpes vulpes flavescens var. cuierascens Birula, .'\nn. Mus. Zool. Acad. Sci. St. 

Petersb. ly: 254. Khorasan, Persia. 
Range: Persia, and Palestine (according to Bodenheimer). 

2 2(j 


VuLPES VULPES pusiLLA Blyth, 1854 

1854. Vulpes pusillus Blyth, J. Asiat. Soc. Bengal, 2j; 729. Salt Range, Punjab. 
1854. Vulpes leucopus Blyth, J. Asiat. Soc. Bengal, 23: 729. Multan, Punjab. 
1875. Vulpes persicus Blanford, Ann. Mag. X.H. 16: 310. Shiraz, Persia. 
Range: North-^Vestern India, from Punjab to Rajputana, Sind, Cutch and Khan- 
desh; Baluchistan, Southern Persia and Iraq. 

Vulpes vulpes griffithi Blyth, 1854 

1854. Vulpes griffithi Blyth, J. Asiat. Soc. Bengal, 23: 730. Kandahar, Afghanistan. 
1845. Vulpes flavescens Hutton, J. Asiat. Soc. Bengal, 14: 344, not of Gray, 1843. 
Range: Afghanistan, Waziristan, Murree in Northern Punjab. 

Vulpes vulpes japonica Gray, 1868 

1868. Vulpes japonica Gray, P.Z.S. 517. Japan. Range includes Hondo, Shikoku and 
Kiushiu, Japan. 

Vulpes vulpes hoole Swinhoe, 1870 

1870. Vulpes hoole Swinhoe, P.Z.S. 631. Near Amoy, Fukien, Southern China. 

1870. Vulpes lineiventer Swinhoe, P.Z.S. 632. Near Amoy, Fukien. 

1907. Vulpes aurantioluteus Matschie, Wiss. Ergebn. Exped. Filchner to China, 10, i : 

168. Tatsienlu, Szechuan, China. 
1923. Vulpes ferrilatus eckloni ]a.cohi, Abh. u. Ber. Mus. f. Tier. u. Volkcrk, Dresden, 

16: 6. Bamutang, three days south-west from Batang, Szechuan, China. Not 

of Przewalsky, 1884. 
Range: Szechuan, eastwards to Fukien in Southern China. 

Vulpes vulpes beringiana Middendorff, 1875 

1875. Canis vulpes var. beringiana Middendorff, Uber Nat. Nord. Ost. Sibir. 4, 2; 

990. Shore of Bering Straits, North-Eastern Siberia. 
1903. Vulpes anadyrensis ]. Allen, Bull. Amer. Mus. N.H. ig: 167. Marcova, Anadyr 

Province, Eastern Siberia. 
191 1. Vulpes kamtschadensis Brass, Aus dem Reich. Pelze, 456. 
1922. Vulpes alopex var. kamtschatica Dybowski, Arch. Tow Nauk. Lwow, /.• 350. 

Kamtchatka, nom. nud. 
Range: North-Eastern Siberia, including Kamtchatka and Anadyr region. 

Vulpes vulpes arabica Thomas, 1902 

1902. Vulpes vulpes arabica Thomas, Ann. Mag. N.H. 10: 489. Muscat, Arabia. 
Ranges south to Aden, north-west to Syria (B.M.). 

Vulpes vulpes alpherakyi Satunin, 1906 

1906. Vulpes alpherakyi Satunin, Isv. Kauk. Mus. 2 {igoj): 46. Geok Tepe, Aralsk 
subdistrict of former Govt, of Elisabetpol, Russian Turkestan. 

Vulpes vulpes kurdistanica Satunin, 1906 

1906. Vulpes kurdistanica Satunin, Isv. Kauk. Mus. 2 (igoj): 48-53. Gelsk Valley, 

Kars district. Western Transcaucasia {probably in extreme North-Eastern 

Asia Minor). 



VuLPES vuLPES icHNUSAE Miller, 1907 

1907. Viilpfs iclmusae Miller, Ann. Mag. N.H. 20: 391. Sarrabus, Sardinia. Range: 
Sardinia, Corsica. 

W'LPES vi-LPES iNDUTA Miller, 1907 

1907. Vulpes indiiliis Miller, .'Vnn. Mag. N.H. 20: 392. Cape Pyla, Cyprus. 

W'LPE.s VULPES siLACEA Miller, 1907 

1907. I'lilpis vulpes silaccus Miller, Ann. Mag. N.H. 20: 393. Near Silos, Burgos, Spain. 

Vulpes vulpes tschiliensis Matschie, 1907 

1907. Vulpes tschiliensis Matschie, Wiss. Ergebn. Filchncr Exped. to China, 70, i: 

1(39. Peiping, Chihli, North-Eastern Clhina. 
(?) 1923. Vulpes huli Sowerby, Nat. in Manchuria, 2: 44. Manchuria. 
Range: Chihli, Shansi, Shensi, Kansu in Northern China, Manchuria? 

\'ULPES VULPES stepensis Brauner, 1914 

1914. Vulpes vulpes stepensis Brauner, Sapiski Novoros ob Estest. //.• 15. {K.V.) Steppes 

near town of Kherson, Russia. Range: Black Sea-Azov steppes, Southern 


A'uLPES VULPES krimeamontana Brauucr, 1914 

1914. Vulpes vulpes knmeamontana Brauner, Sapiski Novoros ob. Estest. //.■ 15-36. 
(N.V.) Mountains of Crimea, Southern Russia. 

Vulpes vulpes caucasica Dinnik, 1914 

1914. Vulpes alopex var. caucasica Dinnik, Svcrikankasa, 2: 449. (N.V.) Near town of 
Vladikawkaz, Caucasus. 

Vulpes vulpes an.\tolica Thomas, 1920 

1920. Vulpes vulpes aualolica Thomas, Ann. Mag. N.H. j: 121. Smyrna, Western Asia 

Vulpes vulpes palaestina Thomas, 1920 

1920. Vulpes vulpes palaestina Thomas, Ann. Mag. N.H. j: 122. Ramleh, near Jaffa, 
Palestine. Range: Palestine and Lebanon. 

Vulpes vulpes jakutensis Ognev, 1923 

1923. Vulpes vulpes jakutensis Ognev, Biol. Mitt. Timiriaz.eiT, /; 116. Taiga south 

from town of Yakutsk, Eastern Siberia. 

Vulpes vulpes diluta Ognev, 1924 

1924. Vulpes vulpes cruagera diluta Ognev, Faun. Voronez Gub. 102-110. Steppe of 

Kamennaia, Bobrov subdistrict of Govt, of Voronej, Russia. A valid race, 
according to Bobrinskii, from the forest-steppe areas of European Russia. 

Vulpes vulpes schrencki Kishida, 1924 

1924. Vulpes vulpes schrencki Kishida, Mon. Jap. Mamm. 47. Sakhalin. Range: to 
Kurile Islands and Hokkaido. 


VuLPES VULPES sPLENDiDissiMA Kishida, 1924 

1924. Vulpes vulpes splendidissima Kishida, Mon. Jap. Mamm. 47. North and Central 
Kurile Islands. 

Vulpes vulpes peculiosa Kishida, 1924 

1924. Vulpes peculiosa Kishida, Chosen. Hanto san no Kitsuna, 4. {N.V.) Korea. 

Vulpes vulpes ochroxantha Ognev, 1926 

1926. Vulpes vulpes ochroxantha Ognev, Ann. Mus. Budapest, 2^: 225. Aksai, Semi- 
rechyia, Eastern Russian Turkestan. 

Vulpes vulpes tobolica Ognev, 1926 

1926. Vulpes vulpes tobolica Ognev, Ann. Mus. Budapest, 25.' 227. Obdorsk, Govt, of 
Tobolsk, Siberia. Range : lower parts of basin of middle and lovver Ob River. 

Vulpes (?) vulpes dolichocrania Ognev, 1926 

1926. Vulpes dolichocrania Ognev, Ann. Mus. Budapest, 2j.' 232. Sidemi, region of 

Southern Ussuri, South-Eastern Siberia. Not listed as a valid form by 

Bobrinskii, 1944. 

Vulpes vulpes alticola Ognev, 1926 

1926. Vulpes vulpes alticola Ognev, Bull. Sci. Inst. E.xplor. Caucas. /.■ 52, 56. Lake 
Gokcha, Transcaucasia (Armenia). 

Vulpes vulpes daurica Ognev, 1931 

1931. Vulpes vulpes daurica Ognev, Mamm. East Europe, 2: 331. Kharangoi, 45 km. 

west from town of Troizkosavsk, Siberia. 
(?) 1922. Vulpes alopex var. ussuriensis Dybowski, Arch. Tow. Nauk. Lwow, /.■ 350, 

nom. nud. 
Range: Amur, Transbaikalia. 

Incertae sedis 

Vulpes alopex var. sibiricus Dybowski, 1922, Arch. Tow. Nauk. Lwow, /.• 350, nom. 

Vulpes kiyomasaiK.i'ihxAa. & Mori, 1929, Lansania, /.• 82, North-Eastern Korea; based 

on a live specimen in Seoul Zoo. 
Vulpes fuliginosus Gray, 1863, Cat. Hodgson Coll. B.NL 6. No locality. 

Vulpes corsac group 

Vulpes corsac Linnaeus, 1 768 Corsac Fox 

Approximate distribution of species: South-Eastern Russia (Kalmuik steppes), 
Volgo-Ural steppes, Russian Turkestan and Kirghizia, to Chinese Turkestan (Zun- 
garia, Bobrinskii), Mongolia, Transbaikalia, and, according to Bobrinskii, Northern 
Manchuria, and Northern Afghanistan. (Blanford quoted it from Persia.) 



V'uLPES CORSAC coRSAC Linnaeus, 1768 

1768. Cariis corsac Linnaeus, Syst. Nat. 12th ed. 3: appendix, 223. Steppes between 

the Ural and Irtish Rivers, Russian Asia. 
1884. Canis ecklom Przcwalski, Reis. Tibet, i i i. Kukunor. 
11)12. Vulpfs corsac nigra Kastschenko, Ann. Mus. St. Petersb. /y; 393. Transbaikalia. 

Not of Borkhausen, 1 797. 
(?) 1944. Viilpes corsac scorodumovi "Dorogostajski, 1935", Bobrinskii, Mamm. 

U.S.S.R. 146 (footnote). Transbaikalia. We are unable to trace an earlier 

reference to this form than that of Bobrinskii, 1944, and that author states 

the form is "of very doubtful reality". 
Range; northern parts of range of the species, Chkalov(=Orcnburg Province), 
Northern Kazakstan, Cis-Altai steppes, Xlongolia, Transbaikalia. 


1035. Vulpi's corsac kalmvkorum Ognev, Mamm. U.S.S.R. 3: 634. Kalmuck Steppe, 
Astrakhan, South-Eastern Russia. 


1935. Vulpes corsac lurkmenicus Ognev, Mamm. U.S.S.R. 3: 635. Turkmen Desert, 
Russian Turkestan. 

Vulpes bengalensis Shaw, 1800 Bengal Fox 

Approxunatc distribution of species: Southern Peninsular India, Travancorc, 
northwards to Sind, Bihar and Orissa, Kangra in Punjab, Haldibari (just south of 
Sikkim), and Nepal. 

Vulpes be.\gale.\sis Shaw, 1800 

1800. Cams bengalensis Shaw, Gen. Zool. /, 2: 330. Bengal. 

1 83 1. Cams kokree Sykes, P.Z.S. loi. Deccan, India. 

1833. Cams ' Vulpes) indicus Hodgson, Asiat. Res. 18, 2: 237. India. Not Canis aureus 

indicus Hodgson, loc. cit. 

1834. Canis (Vulpes) rufescens Gray, Hardwicke's 111. Ind. Zool. 2, pi. 3. India. 
1837. Canis chrysurus Gray, Charlesw. Mag. N.H. /.• 577. Nepal. 

1837. Vulpes hodgsonii Gray, Charlesw. Mag. N.H. /.• 578. Nepal. 

1838. Vulpes xanthura Gray, P.Z.S. 183J: 68. Nepal. 

Vulpes riippelli Schinz, 1825 Sand Fox 

Appniximatc distribution of species: Sudan, Somaliland, Asbcn, north to Southern 
Algeria, Libva and Egypt; Sinai, Southern Arabia; Persian Baluchistan and 

Vulpes ruppllli ruppelli Schinz, 1825 

1825. Cann ruppelli (sic) Schinz, Cuviers Thierreich, ./.• 508. Dongola, Sudan. 

182(3. Cams fa/mlicus Cretzschmar, in Ruppell, Atlas zu d. Reise im nordl. Afrika, 

Saugclh. 15. Nubian Desert and Kordofan. Ranges north to Egypt 

( Fliiwerj. 



191 2. Vulpes [Megalotis) famelicus zarudnyi Birula, Ann. Mus. Zool. Sci. St. Petersb. 
ly: 270. Kala-i-bid, Prov. Makran (Sargad), Persian Baluchistan. Ranges 
into Afghanistan (B.\L). 

Vulpes ruppelli caesia Thomas & Hinton, 192 1 

1 92 1. Vulpes riippellii caesia Thomas & Hinton, Nov. Zool. 28: 5. Southern side Mt. 

Baguezan, Asben, Western Sahara. Ranges north to Ahaggar, Southern 


Vulpes rijppelli cyrenaica Festa, 1921 

1 92 1. Vulpes cyrenaica Festa, Boll. Mus. Zool. Anat. Comp. Univ. Torino, j^, 740: 3. 
Near Benghazi, Cyrenaica, Libya. 

Vulpes ruppelli sabaea Pocock, 1934 

1934. Vulpes ruppelli sabaea Pocock, Ann. Mag. N.H. i^: 636. Rub al Khali, Arabia. 

Vulpes pallida group 

Vxilpes cana Blanford, 1877 Blanford's Fox 

Approximate distribution of species: Kopet Dag, in South-^Vestern Russian 
Turkestan; Afghanistan, North-Eastern Persia, Baluchistan. 

Vulpes cana Blanford, 1877 

1877. Vulpes canus Blanford, J. Asiat. Soc. Bengal, 46, 2: 321. Gwadar, Baluchistan. 
1907. Vulpes cana var. nigricans Shitkow, Zool. Anz. 52.' 448. Bokhara, Russian 

Vulpes ferrilata group 

Vulpes ferrilata Hodgson, 1842 Tibetan Sand Fox 

Approximate distribution of species: Tibet and Nepal. 

Vulpes ferrilata Hodgson, 1842 

1842. Vulpes ferrilatus Hodgson, J. Asiat. Soc. Bengal, //.• 278. Near Lhasa, Tibet. 

Genus FENNECUS Desmarest, 1804 

1804. Fennecus Desmarest, Diet. d'Hist. Nat. 24, Tabl. meth. Mamm. 18. Fennecus 

arabicus Desmarest = Canis zcda Zimmermann. 
181 1. Alegalotis lUiger, Prodr. Syst. Mamm. et Avium, 131. Canis cerda Gmelin = 

Canis zerda Zimmermann. 

I species: Fennecus z^rda, page 231 

Fennecus zerda Zimmermann, 1 780 Fennec Fox 

Approximate distribution of species: Morocco, Algeria, Libya, Egypt, thence to 
Sinai and Arabia, south to the Sudan and Asben. 



Ff.n' zerda Zimmcrmann, 1 780 

I 780. Cams zerda Zimniermaiiii, Geogr. Ges. 2: 247. Sahara, and other parts iii'Xnrth 

Africa behind the Atlas. 
1777. Vulpcs minimus saarcnsis Skjoldebrand, K. S\cnska \'et. Akad. Hand!. Sti»-k- 

holm, j<?.- 267. "This name if considered vahd would supersede Cams .yrda, 

but although the author states that he wishes to include the animal in the 

Liiinean system, he gives a trinomial name" (Glover Allen'i. Algerian 

1788. Canis cerdo Gmclin, Linn. Syst. Nat. 13th ed. /.• 73. Sahara. 
179'^. Viverra aurila Meyer, Zool. Entdeck. in Xeu Holland u. Africa, 91. Biskra, 

Beni Mezzab and Weryleh, Algeria. 
1804. Femiecus arabicus Dcsmarest, Diet. H.X. 24. Tabl. mcth. Manmi. 18. "Barbary, 

Nubia, Ab)ssinia." 
181 I. Mt-nalotis cerda Illiger, Prodr. Syst. Mamm. 131. 
1820. Femiecus bnicei Desmarcst, Encyclop. Meth. Mamm. 235. Libya, Tunis, Algeria, 

1827. Cams frmieciis Lesson, .\Lmuel .\Limm. ib8. 
1842. Vidpes denhamii Boitard, Le Jardin des Plantes, 213. "L-itcrior of Africa." 

Genus NYCTEREUTES Temminck, 1839 

1839. Avcleiriilcs Temminck, in Wan der Hoevens Tijdschr. Nat. Ges. Phys. j; 285. 
Mrctcieules z-ivernmis Temminck. 

I species: Nvcteri-ules procvonoides, page 232 

Nyctereutes procyonoides Gray, 1834 Raccoon-Dog 

Approximate distribution of species: Amur and Ussuri region of Eastern Siberia; 
Japan, Manchuria, states of Shansi, Szechuan, Yunnan, south-eastwards to Fukien 
and district, in C'.hina; Tonkin, in Northern Indo-Ghina. 

Nyctereltes PROCA'ONOiDEs PROCYONOIDES Gray, 1834 

1834. Cams procyonoides Gray, Illustr. Ind. Zool. 2: pi. i. Vicinity of Canton, Southern 

China" {see G. Allen, 1938, .\Limm. China & Mongolia, /.• 346). 
1904. Nvctereutes sinensis Brass, Nutzbare Tiere Ostasiens, 22. Yangtze \'allcy, China. 
1907. S'vctereutei stegmanni Matschie, Wiss. Ergebn. Filchners Exped. to China, 10, i : 

175, i8u. ifising-an-fu, Chinkiang, Kiangsu, Southern East China. 
Ranc^e: Clhinese range of the species, except Yunnan. Tonkin, in Indo-China. 

Nyctereutes procyonoides viverrinus Temminck, 1844 

1844. .Xrclereiites viverrinus Temminck, Siebolds Fauna Japonica, .\Limm. 40, pi. 8. 

(?J 1904. .\vctcrcules albus Beard, Scientific American, c)i: 287. "Based on a white 

specimen in the New York Zoological Park, said to be from Hokkaido, 

Japan." But listed as a \alid race for Hokkaido by Kuroda, 1938, .\Limm. 

Range includes also Hondo, Shikoku, Kiushiu. 



Nyctereutes procyonoides ussuriensis Matschie, 1907 

1907. Nyctereutes ussuriensis Matschie, Wiss. Ergebn. Filchners Exped. to China, 10, 

1 : 1 78. Near mouth of Ussuri River, Eastern Siberia. 
1907. Nyctereutes amurensis Matschie, loc. cit. 179. Amur. 

Nyctereutes procyonoides koreensis Mori, 1922 

1922. Nyctereutes koreensis Mori, Ann. Mag. N.H. 10: 607. Giseifu, near Seoul, Korea. 

Nyctereutes procyonoides orestes Thomas, 1923 

1923. Nyctereutes procronoides orestes Thomas, Ann. Mag. N.H. //.• 657. North-western 

flank Likiang Range, Yunnan, about 10,000-12,000 ft., China. 

Genus CUON Hodgson, 1838 

1838. Cuon Hodgson, Ann. Mag. N.H. /; 152. Canis primaevus Hodgson. 

1839. Chrysaeus H. Smith, Jardine's Nat. Libr. Mamm. 25.- 167. Canis dukhunensis 

1888. Cyan Blanford, Fauna Brit. India, Mamm. /.• 142. (Emendation of Cuon.) 
1888. Anurocyon Heude, Mem. H.N. Emp. Chin. 2: 102. Anurocyon clamitans Heude = 

Canis lepturus Heude. 

Pocock recognized only one species in this genus, for which the earliest name is 
C. alpinus Pallas, 181 1. 

I species: Cuon alpinus, page 233 

Cuon alpinus Pallas, 181 1 Dhole, Red Dog, or Indian Wild Dog 

Approximate distribution of species: Eastern Russian Turkestan (Eastern Pamirs, 
Tianshan and Tarbagatai Mountains), Russian Altai, .Southern Cisbaikal and 
Southern Transbaikal, Amur and Ussuri regions of Eastern Siberia; Chinese 
Turkestan, part, according to Ognev, probably Southern Tibet, Korea, Sakhalin; 
states of Szechuan, Yunnan, Fukien, in China; Peninsula of India, from Coorg and 
Nilgiri Hills northwards to Kashmir, thence to Nepal, Burma, Tenasserim; Indo- 
China, Malay States, Sumatra and Java. 

Cuon alpinus alpinus Pallas, 181 1 

181 1. Canis alpinus Pallas, Zoogr. Ross. Asiat. /.• 34. Near Udskoi Ostrog, Amurland. 
Range includes Manchuria and Sakhalin. 

Cuon alpinus dukhunensis Sykes, 1831 

1 83 1. Canis dukhunensis Sykes, P.Z.S. 100. Deccan, Peninsular India. Range: India, 
south of the Ganges. 

Cuon alpinus primaevus Hodgson, 1833 

1833. Canis primaevus Hodgson, Asiat. Res. 18, 2: 221. Nepal. 

1863. Cuon grayiformis Hodgson, in Gray, Cat. Hodgsons Coll. B.M. 2nd ed. 5. 

Range: Kumaon, Nepal, Sikkim, Bhutan. 

Note: Osgood (1932), On Indo-Chinese Mammals, Field Mus. N.H.^ool. 18: 193, 



et seq., uses for the Wild Dogs of Indo-China the name Cuon rutilans Muller, 1839, 
Temm. Verh. nat. ges. J\^ed. oveiz- bezitt. -Zool. 27, 51, which according to Chasen, 1940, 
Handlist Malaysian Mamm., is a synonym of Cuon alpinusjavanicus Desmarest, 1820, and 
came from Java. These Indo-Chinese \Vild Dogs are now referred to C. a. aduslus 
Pocock (below). 

Cuon .•vlpinus lepturus Heude, 1892 

1892. Cuon lepturus Heude, Mem. H.N. Emp. Chin. 2. 2 (footnote), 102. Poyang 

Lake, south of the Yangtze, Kiangsi, Clhina. 
1892. Amirocxon clamitarn Heude, loc. cit. Taihu, near mouth of the Yangtze, Clhina. 

Cuox .NLPiNUS HESPF.Rius Afauasiev & Zolotarev, 1935 

1935. Cyan alpinus hespenus Afanasiev & Zolotarev, Bull. Acad. Sci. U.S.S.R. 7: 427. 

Aksai district of Semiryechensk region, Eastern Russian Turkestan. 
(?) 1936. Cuon javanicus jason Pocock, P.Z.S. 51. Altai Mountains. 

Cuon alpinus infuscus Pocock, 1936 

1936. Cuon javanicus infuscus Pocock, P.Z.S. 38, fig. la. Moulmein, Tena.sserim. 

Cuon alpinus fumosus Pocock, 1936 

1936. Cuon javanicus fumosus Pocock, P.Z.S. 49. Western Szechuan, China. 

Cuon alpinus laniger Pocock, 1936 

1936. Cuon javanicus lanigcr Pocock, P.Z.S. 50. Kashmir. Probably ranges to Lhasa, 
Southern Tibet. 

Cuon alpinus adustus Pocock, 1941 

1 941. Cuon alpinus adustus Pocock, Fauna Brit. India, -.• 156. Upper Burma. Range: 
Upper Burma, Indo-China. 

Genus LYCAON Brookes, 1827 

1827. Lycaon Brookes, in Griffith Ckiv. Anim. Kingd. 5.- 151. Lycaon tricolor Brookes = 

Hyaena picta Temminck. 
1829. Cyn'hyaena F. Cuvier, Diet, des Sci. Nat. 59; 454. Hyaena picta Temminck. 
1842. Hycnoidcs Boitard, Le Jardin des Plantcs, 215. Hyaena picta Temminck. 

I species: Lycaon pictus, page 234 

Lycaon pictus Temminck, 1820 African Hunting Dog 

Approximate distribution of species: Africa, from South-West Africa and Kruger 
National Park (Transvaal), northwards to Somaliland and the Sudan, Lake Chad 
district, Dahomey, and (apparently) Southern Algeria. 

(Lycaon pictus pictus Temminck, 1820. Extralimital) 

1820. Hyaena picta Temminck, Ann. Gen. Sci. Phys. j: 54, pi. 35. Coast of Mozam- 



Lycaon pictus sharicus Thomas & Wroughton, 1907 

1907. Lycaon pictus sharicus Thomas & Wroughton, Ann. Mag. N.H. ig: 375. Mani, 

Lower Shari River, east of Lake Chad (French Congo). 
1 915. Lycaon ebermaieri Matschie, S.B. Ges. Nat. Fr. Berlin, 369. Lake Chad region. 

G. Allen, 1939, Checklist African Mammals, 191, quotes the race L. p. 

sharicus from Tanezrouft, Algeria. 


Genera: Helarctos, page 241 
Alelursus, page 241 
Selenarctos, page 239 
Thalarctos, page 240 
Ursus, page 235 

See particularly Pocock, 1932, The Black and Brown Bears of Europe and Asia, 
J. Bombay N.H. Soc. 35, i : 771 ; and J. Bombay KH. Soc. 36,2: loi . In this paper, a 
key to all genera listed above, except Thalarctos, will be found. Miller (1912, 285) 
gives the generic characters oi Thalarctos. Simpson (1945, 225) follows Pocock in his 
classification of the bears, and we entirely agree with his remarks on the species and 
genera. It must be admitted, however, that Selenarctos might be considered a sub- 
genus oi Ursus, and it may be noted that Bobrinskii (1944) refers all Russian bears, 
including Thalarctos, to the genus Ursus, in which fp. 136) he lists three subgenera. 
Pocock (1941, 169) gives a short note on Thalarctos compared with the four British- 
Indian genera, and comparison of Miller's figures of skulls of Thalarctos and Ursus 
with Pocock's figures of the skulls of the other three genera enables Thalarctos to be 
quite easily distinguished by skull alone, apart from its somewhat unique external 
appearance. Each of the genera listed here contains one species only in the present 

Genus URSUS Linnaeus, 1 758 

1758. Ursus Linnaeus, Syst. Nat. loth ed. /.• 47. Ursus arctos Linnaeus. 

1864. Euarctos Gray, P.Z.S. 692. Ursus americanus Pallas. Valid as a subgenus. 

1864. .Myrmarclos Gray, P.Z.S. 694. Myrmarctos eversmanni Gray = Ursus arctos 

1898. Ursarctos Heude, Mem. H. N. Emp. Chin. 4, i : 17 (yesoensis). 
1898. Melanarctos Heude, Mem. H.X. Emp. Chin. 4, i: 18. Melanarctos cavifrons 

Heude = Ursus lasiotus Gray. 
1923. Mylarctos Lonnberg, P.Z.S. 91. Ursus pruinosus Blyth. 

I species in Eurasia : 

Ursus arctos, page 236 

Q 235 


Ursus arctos Linnaeus, 1758 Brown Bear 

Approximate distribution of species: Sweden, Norway, Finland, Estonia, Poland, 
Ci-echoslovakia, Austria, Yugoslavia, Albania, Rumania, Bulgaria, Greece, Italy 
(Abruzzi and Trentino), France (Pyrenees and, doubtfully, in Forest of Vcrcors, 
(Drome), Spain (Pyrenees and Asturias). Most of the U.S.S.R.; according to Bobrin- 
skii, "whole of the forest zone, whence it penetrates in the summer far into the 
lundra, Karaginskii Island in Bering Sea, the Shantar Islands, Sakhalin. Mountains 
ol' Central Asia, all the mountainous parts of the Caucasus; does not occur in Crimea". 
Mongolia, Manchuria, Japan; Tibet, Kansu, probably Szechuan. Syria (extinct in 
Palestine), Persia, Asia Minor. Kashmir, Punjab. Also in North America. 

Ursus arctos arctos Linnaeus, 1758 

1758. Ursus arctos Linnaeus, Syst. Nat. loth ed. /.• 47. Sweden. 

1772. Ursus ursus Boddaert, Kortbegrip van het zamcnstel der Nat. /.• 46. (jV.F.) 

Renaming of arctos. 
1788. Ursus arctos niger Gmelin, Syst. Nat. 13th ed. /.■ 100. Northern Europe. 
1788. Ursus arctos fuscus Gmelin, loc. cil. Alps. 
1 788. Ursus arctos albus Gmelin, loc. cil. Locality unknown. 
1792. Ursus arctos griseus Kerr, Anim. Kingd. 184. Germany. 

1797. Ursus arctos rufus Borkhausen, Deutsche Fauna, /.■ 46. Swiss and Tirolean Alps. 

1798. Ursus hadius .Schrank, Fauna Boica, /.• 55. Forests on Bohemian boundary. 

1 808. Ursus fuscus Tiedemann, Zool. /.• 374. Substitute for arctos ; not of Gmelin, 1 788. 

1814. Ursus alpinus Fischer, Zoognosia, ^.- 161. .''Alps. 

1820. Ursus arctos major Nilsson, Skand. Fauna, /; 112. Southern Scandinavia. 

1820. Ursus arctos minor Nilsson, loc. cit. 123. Northernmost Scandinavia. 

1827. Ursus arctos brumieus Billberg, Synop. Faunae Scandinaviae, 15. Northern 


1827. Ursus arctos annulatus Billberg, loc. cit. 15. Northern Scandina\ia. 

1827. Ursus arctos argenteus Billberg, loc. cit. 15. NVirthern Scandinavia. 

1827. Ursus arctos mvnncphagus Billberg, loc. cit. iG. Northern Scandinavia. 

1828. Ursus formicarius Billberg, Syncips. Faun. Scand. 2nd ed. 16. Renaming of 


1829. Ursus pvrenaicus Fischer, Synops. Mamm. 142. Asturias, Spain. 
1829. Ursus norvegicus Fischer, loc. cit. Norway. 

(?) 1836. Ursus falciger Reichenbach, Regn. Anim. Icon. /.■ 32. Pyrenees. (N.V.) 
("afterwards supposed to be an individual of'U.fero.x' See Naturgesch. des 
In-und Auslands, Raubsaugeth. p. 299, 1852" as quoted by Miller, 191 2 Cat. 
Mamm. W. Europe, 286. (U.fcro.x = U. horrihilis Ord, from North America.) 

1840. Ursus cadaverimts Evcrsmann, Bull. Soc. Imp. Nat. Moscow, 11. Renaming of 
U. arctos. 

1840. Ursus Iniigiroitrii E\ersmann, Inc. cil. Renaming nf formicarius. 

(?) 1847. Ursus euryrhimis Nilsson, Skand. Fauna, 2nd ed. /.■ 212. ? Sweden. (Type an 
individual raised in captivity.) 

l8-,5. Ur\in arctos aureus Fitzinger, W'iss. pop. Nat. der Saugcth. /.■ 372. Eastern 

1864. Ursus arctos var. (i 1 normalis Gray, P.Z.S. 682. (Renaming c^i arctos.) 

1864. Ursus arctos sub-var. (a) scandinaricus Gray, P.Z.S. 682. 

i!i()4. Ursus arctos sub-var. (c) rossicus Gray, P.Z.S. 682, iiom. mid. 



1864. Ursus arclos sub-var. (f) polonicus Gray, P.Z.S. 682. Poland. 

1864. Ursus arctos var. (2) grandis Gray, P.Z.S. 684. "North of Europe," a male 

purchased at Hull, living in the Zoological Gardens from 1852 to 1863. 
1864. Ursus arclos var. (4) stenorostris Gray, P.Z.S. 685. Poland, based on Cuvier, 

1823, Oss. Fossiles, 4: 332, 2nd var. 
1864. Myrinarctos eversmanni Gray, P.Z.S. 695. Norway. 
(?) 1905. Ursus formicarius (Eversmann) Bieler, C.R. Sixieme Congres Internal, de 

Zool. Berne, 248. Switzerland. 
(?) 1 92 1. Ursus arctos marsicanus Altobello, Fauna Abruzzo e Molise, .Mamm. 15. 

Abruzzo, Italy. 
Range: European range of the species, eastwards as far as the Stanovoi Range, 

Ursus .\rctos collaris Cuvier & Geoffrey, 1824 

1824. Ursus collaris Cu\ier & Geoffroy, H.N. Mamm. pt. 42, pi. 212. Siberia. 

1864. Ursus arctos var. sibiricus Gray, P.Z..S. 682. Siberia. 

1924. Ursus arctos jeniseensis Ognev, Nature & Sport in Ukraine, /, 2: iio. River 
Ungut, taiga in mountains in surroundings of Krasnoiarsk, Ycnessei Pro- 
vince, Siberia. 

This name is not used by the Russian authors Ognev and Bobrinskii, but the name 
appears to be the second valid name in the Palaearctic for the species, and is retained 
by Pocock, 1932, J. Bombay N.H. Soc. 55, 4: 793. 

Ursus arctos isabellinus Horsfield, 1826 Red Bear 

1826. Ursus isabellinus Horsfield, Trans. Linn. Soc. Zool. /j.- 334. Mountains of 

1873. Ursus leuconyx Severtzov, Mem. Soc. Amis. Sci. Nat. Mosc. 8: 79. Upper part 

of valley of River Naryn, Tianshan Mountains. 

1 924. Ursus pamirensis Ognev, Nature & Sport in Ukraine, 7,2: 1 1 1 . Pamir Mountains. 
Range: Tianshan, Pamirs, Afghanistan? W'aziristan, Kashmir, Punjab. Bobrinskii 

(1944) lists leuconyx as a valid form, but Pocock (1932, 1941) states it is the same as 
isabellinus, which has priority. 

Ursus arctos svriacus Hemprich & Ehrenberg, 1828 

1828. Ursus svriacus Hemprich & Ehrenberg, Symb. Phys. /.• sig. a, pi. i. Near \illagc 

of Bischerre, Mt. Makmel, Lebanon. 
191 7. Ursus schmitzi Matschie, S.B. Ges. Nat. Fr. Berlin, 33. Mt. Hermon, Palestine. 

In addition, Pocock appears to treat the following names as synonyms: 
1 85 1. Ursus arctos var. meridionalis Middendorff, \'erh. Russ. Kais. Min. Ges. 80. 

1913. Ursus arctos lasistanicus Satunin, Tr. Obshch. Chernomorsk poberezh'ya, 2: 27. 

Black Sea coast. 
1919. Ursus arctos var. caucasicus Smirno\-, Bull. .\Ius. Cauc. 12: iij. Pasanaur, 

southern slope of Central Caucasus mountains. 
1919. Ursus arctos arctos natio dinniki Smirnov, Bull. Mus. Cauc. 12: 122. Chatakh 

Borchalinsk subdistrict of Govt, of Tiflis, Caucasus. 

1925. Ursus arctos smirnovi Lonnberg, Fauna och Flora, /.• 28. Northern slopes of 

main chain of Caucasus. 



Ursus arctos syr[aclis [contd.] 

1925. Ursus arctos persicus Lonnberg, Fauna och Flora, /; 28. Mazanderan, Northern 

Range; Syria, Asia Minor, Persia and the Caucasus. It should be noted that Bobrin- 

skii lists two races of this species from the Caucasus, syriacus (South-Western 

Transcaucasia! and caucasicus (other parts of the Caucasus). 

Ursus .•\rctos beringi.^nus Middendorff, 1853 

1853. Ursus arctos var. beringiana Middendorff, Sibir. Reise, ;■, 2 : 4, pi. i, figs. 1-6. 

Great Shantar Island, Sea of Okhotsk. 
1855. Ursus piscator Pjjcheran, Rev. Mag. Zool. y: 392. Petropaulovski, Southern 

(?) i8q8. Ursus mandchuncus Heude, Mem. H.N. Nat.Emp. Chin. 4: 23-24, pi. 7, 

figs. i-K. Near Vladivostock. Bobrinskii thinks that this should probably 

stand as a valid race, but Pocock synonymized it. 
1924. Ursus arctos kolvmensis Ognev, Nature & Sport in Ukraine, /, 2 : 112. Saborzevo, 

River Kolyma, north-west of Sea of Okhotsk. 
Range : Siberia, east of Stanovoi Range, particularly in Kamtchatka, Ussuri and 

Ursus arctos pruixosus Blyth, 1854 Blue Bear 

1854. Ursus pruinosus Blyth, J. Asiat. Soc. Bengal, 22: 589. Lhasa, Tibet. 

1883. Ursus lagornviarius Przewaiski, Third Journ. in Cent. Asia, 216. Kuku-Shili 

Range, 35° N., 92' E., Tibet. 
Range: Tibet, Kansu. 

Ursus arctos lasiotus Gray, 1867 

1867. Ursus lasiotus Gray, Ann. Mag. N.H. 20: 301. Interior of Northern China. 

1844. Ursus fcrox Tcmmmck, Fauna Japonica, 29, not of Rafinesque, 1817. 

1897. Ursus arctus yesoensis Lydekker, P.Z.S. 422. Yeso (= Hokkaido), Japan. 

1898. Ursus melanarctos Heude, Mem. H.N. Emp. Chin. 4: 17. Yeso (= Hokkaido), 

1901. Melanarctos cavifrons Heude, Mem. H.N. Emp. Chin. 5, i: i. Tci-tci-tar 

(Tsitsihar), North-\Vestern Manchuria. 
(?) 1924. Ursus arctos baikalcnsis Ognev, Nature & Sport in Ukraine, /, 2: 112. 

Province of Irkutsk, near Lake Baikal, Eastern .Siberia. 
Range: Mongolia, Manchuria, Hokkaido and Kurilc Islands, Korea. 
The following races may be of doubtful status : 

Ursus arctos crowt/ieri Schinz, 1844, Synops. Mammalium, /.■ 302 (based upon 
the "Bear of Mount .\tlas" of Blyth, 1841, P.Z.S. 65. "Foot of the Tetuan 
mountains, about twcnty-fi\c miles from that of the Atlas." Doubts have 
been thrown on the existence of this bear, but though it is now e.xtinct it 
almost certainly did exist in 1844 (see Harper, 1945, Extinct and Vanishing 
Mammals of the Old World, 230) ). 
Ursui arctos shanorum Thomas, 1906, Abstr. P.Z.S. 17. Said to be from the Shan 
.States, LIppcr Burma, where the species probably does not occur. .See 
Pocock ' 1 94 1, 18-)). 
f V.H/s (if(tn\ finsnu-riiis Bolkay, 1 925, Nov. Mus. .Saraje\'o, /.• 8. Bosnia, Yugoslavia. 



Genus SELENARCTOS Heude, 1901 

1 90 1. Selenarctos Heude, Mem. H.N. Emp. Chin, j.- 2. Ursus thibetanus Cuvier. 
1917. Arcticonus Pocock, Ann. Mag. N.H. 20: 129. Ursus thibetanus Cuvier. 
1938. Euarctos G. Allen, Mamm. China & Mongolia, /.• 330 (in part); not Euarctos 
Gray, 1864. 
I species: Selenarctos thibetanus, page 239 

Selenarctos thibetanus G. Cuvier, 1823 Asiatic Black Bear 

Approximate distribution of species: Amur and Ussuri regions of far east of 
Siberia; Japan, Manchuria, Formosa; most of China, westwards to Kansu and 
Szechuan, south to Fukien and Hainan; Indo-China, Siam; from Burma and Assam 
westwards to Nepal, Kashmir and Baluchistan; Afghanistan (Bobrinskii) . 

Selenarctos thibetanus thibetanus G. Cuvier, 1823 
1823. Ursus thibetanus G. Cuvier, Ossements Foss. 4: 325. Sylhet, Assam. 
1841. Ursus torquatus Wagner, in Schreb. Saugeth. Suppl. 2: 144. Renaming of 

1876. Ursus sp. (? Melursus labiatus) Blanford, E. Persia, 47. Not of Blainville, 181 7. 
Range: from Nepal eastwards through Assam, Burma, and Siam to Annam. 

Selenarctos thibetanus japonicus Schlegel, 1857 

1857. Ursus japonicus Schlegel, Handl. Dierkunde, /.• 42. (Assumed to be) Japan. 

1897. Ursus rexi Matschie, S.B. Ges. Nat. Fr. Berlin, 72. Japan. 

Range: Hondo, Kiushu, PShikoku. 

Selenarctos thibetanus formosanus Swinhoe, 1864 

1864. Ursus formosanus Swinhoe, P.Z.S. 380. Formosa. 

(?) 1922. Selenarctos melli Matschie, Arch. Nat. 88, 10: 34. Hainan. 

Pocock thought this was either a synonym oi formosanus or the typical race. G. 
Allen (1938) listed it as a valid race from Fukien and Hainan. 

Selenarctos thibetanus gedrosianus Blanford, 1877 

1877. Ursus gedrosianus Blanford, Proc. Asiat. Soc. Bengal, 204. Tump, 70 miles north 

of Gwadar, on the Mekran coast, Baluchistan. 

Selenarctos thibetanus ussuricus Heude, 1901 

1901. Selenarctos ussuricus Heude, Mem. H.N. Emp. Chin. 5, i : 2, pi. ii, fig. 10. 

Ussuri region, Eastern Siberia. 
1928. Selenarctos thibetanus wulsini Howell, Proc. Biol. Soc. Washington, 41: 115. 

Eastern Tombs, Chihli, North-Eastern China. 
Range: Northern China, Manchuria, Amurland and Ussuri, Korea. 

Selenarctos thibetanus mupinensis Heude, 1901 

1901. Selenarctos mupinensis Heude, Mem. H.N. Emp. Chin. 5, i : 2, pi. ii, figs, i, 2, 9. 

Moupin, Szechuan, China. 
1901. Selenarctos leuconyx Heude, loc. cit., figs. 3, 4, 8. Taipei Shan, South- Western 

Shensi, China. 




loot), i'rsiis tonjuatits macthiUi Lydekker, P.Z.S. 609. Tatsienlu, Szechuan, China. 
11^-20. Visits clarki Sowerby, J. Mamm. /.■ 226. New name for leuconyx Hcude. A 

svnonvm oi'/hihf tenuis according; to G. Allen 1 1 938) , but a valid race according 

to Pocock. 
Rancjc includes Shcnsi, .Szxchuan and Hupch, China. 

Sele.\.a,rctos'us lamger Pocock, 1932 

1932. Selenarctos thihetanus laniger Pocock, J. Bombay N.H. Soc. :::G: i 15. Aru, Upper 

Lidder Valley, Kashmir. 
I?) 1864. Ursus torqualiis var. arhoreus Gray, P.Z.S. 688. Darjeeling. 
Range: Kashihir and Punjab. 

Genus THALARCTOS Gray, 1825 

1825. Thalanios Gray, Ann. Philosophy, X.S. 10: 62, July 1825. Thalarctos polaris 

Gray = Ursus maritimiis Phipps. 
1825. Thalassarctos Gray, Ann. Philosophy, N.S. 10: 339. November 1825. 
1896. Thalassiarchus Kobelt, Bericht Senckenbcrg. naturf. Ges. Frankfurt am Main, 

93. (Substitute for Thalarclos.) 

I species : Thalarctos maritimiis, page 240 

Thalarctos maritimus Phipps, 1774 Polar Bear 

Approximate distribution of species: Arctic regions of Europe, Asia and North 
America. "South on floating ice occasionally to the northern coast of Norway" 
(Miller). In U.S.S.R., "only occurs by chance on European coasts. Does not penetrate 
far on to the mainland. Numerous on sea coasts of the Kara, Laptev, Eastern Siberian 
Seas and Clhukotskaya Seas, and on Dixon Island, the Novosibirskie Islands, Med- 
\czhie Islands and Wrangel Island. It is rare on the west coast of the south island of 
Novaya Zemlya, and common on the northern island and in Spitzbergen. On 
\'aigach and Kolguev Islands it is very rare and occurs only in winter. Cases are 
known of its having been carried on icefloes in the winter to the Murman coast and 
Kanin Peninsula. In Bering Sea it is already rare and on the Anadyr coast only 
occurs in exceptional cases. It is not known to occur in Kamtchatka, but has several 
times been carried on icefloes to Sakhalin and was once observed in the north of the 
Sea of Okhotsk ' Tui Bay)". According to Kuroda's list (IQ38J has been recorded 
from Japan iKiniles, Hokkaido, ? Hondo;. 

Th.\larctos m.\riti.\ius maritimus Phipps, 1774 

1774. Ursus maritimus Phipps, \'oyage toward North Pole, 18",. Spitzbergen. 

(?) 1776. Ursus marinus Pallas, Reise durch verschiedenc Pro\inzcn dcs Russ, Reichs, 

J.' 691. Arctic Ocean, Siberia. 
1792. Ursus polaris .Shaw, Mus. Leverianum, /.■ 7. Renaming ot marinus. 
'') 1908. Thalassarctos jeriaaisis Knottnerus-Mayer, S.B. Ges. Nat. Fr. Berlin, 184. 

Jena Island, Spitzbergen. 
?J 1908. Thalassarctos spi/rhcrffcrisi', Knottncrus-Mascr, loc. cit. .Seven, 



Genus HELARCTOS Horsfield, 1825 
1825. Helarctos Horsfield, J. Zool. 2: 221, 233. Ursus malayanus Raffles. 
I species: Helarclos malayanus, page 241 

Helarctos malayanus Rafiles, 1821 Malayan Sun Bear 

Approximate distribution of species: Burma, Indo-China, Siam, Malay Peninsula, 
Sumatra, Borneo. Possibly, but not certainly, from Szechuan or some adjacent 
region of Southern China. 

Helarctos malayanus malayanus Raffles, 1821 

1 82 1. Ursus malayanus Raffles, Trans. Linn. Soc. London, /5:254.Bencoolcn, Sumatra. 
1901. Helarctos annamiticus Heude, Mem. H.N. Emp. Chin. 5, i.- i, pi. i, figs. 1-2. 

Annam, Indo-China. 
1906. Ursus malayanus wardi Lydekker, P.Z.S. 999. Thought to be from Tibet (or 

Szechuan or Yunnan, G. Allen). 
Range: as in the species, except Borneo. 

Genus MELURSUS Meyer, 1793 

1793. Melursus Meyer, Zool. Entdeck. 155. Bradypus ursinus Shaw. 

1809. Arceus Goldfuss, Verh. Nat. Saug. 301. Bradypus ursinus Shaw. 

181 1. Prochilus Illiger, Prodr. Syst. Mamm. log. Bradypus ursinus Shaw. 

1814. Chondrorhy7ichus Fischer, Zoogr. 2- H^- Bradypus ursinus Shaw. 

I species: Melursus ursinus, page 241 

Melursus ursinus Shaw, 1791 Sloth Bear 

Approximate distribution of species: Ceylon, Southern Peninsula of India, north- 
wards to Central Provinces, Bihar, Bengal and Assam. ? Darjeeling. 

Melursus ursinus ursinus Shaw, 1791 

1 791. Bradypus ursinus Shaw, Nat. Misc. 2 (unpaged), pis. 58-59. Patna, on the 

Ganges, Bengal. 
1793. Melursus lybius Meyer, Zool. Entdeckung. 156. "Africa interior." 
1809. Arceus niger Goldfuss, Verh. Nat. Saug. 301 (teste Palmer). 
1817. Ursus labiatus Blainville, Bull. Soc. Philom. 74. 
1820. Ursus longirostris Tiedemann, Abhandl. Bar. Faulthier, i i. 
Range: as above, Ceylon excluded. 

Melursus ursinus inornatus Pucheran, 1855 

1855. Melursus inornatus Pucheran, Rev. Mag. Zool. j: 392. Ceylon. 




Genera: Ailuropoda, page 242 
Ail III us, page 242 
Simpson (1945, 226) refers the Asiatic Pandas to this (principally American) 
family, as a subfamilv, the Ailurinae. Pocock referred the two to two distinct fiimilies, 
Ailuridae and Ailuropodidae. G. Allen (1938) referred Ailiirus to the Procyonidae, 
but retained the family Ailuropodidae. While the differences between the two genera 
seem very wide, we follow Simpson, preferring his classification to the very split one 
offered by Pocock for the Raccoons and their allies. 

Subfamily Ailurinae 

Genus AILURUS Cuvier, 1825 

1825. Ailurus F. Cuvier, in E. Geoffroy & Cuvier, H.N. Mamm. 5 (50), 3. Ailurus 

fulgens Cuvier. 
1841. Arcladurus Gloger, Gcmein. Hand. Nat. /.■ xxviii. A. fulgens. 

1846. Adurus Agassiz, Nomcnclator Zool. index, Univ. 9. Emend, pro Ailurus Cuvier. 

I species; Ailurus fulgnis, page 242 

Ailurus fulgens F. C;uvier, 1825 Red Panda 

Approximate distribution of species: ^'unnan and Szechuan, in China; Northern 
Burma, Sikkim and Nepal. 

Ailurus fulgens fulgens F. Cuvier, 1825 

1825. Ailurus fulgens Cuvier, in Geoffroy & Cuvier, H.N. Manmi. jj (50): Panda, 3. 
Locality unknown ("East Indies"). 

1847. Ailurus ochraceus Hodgson, J. Asiat. Soc. Bengal, 16: 1 1 18. "Sub Himalayas," 

from 7,000 to 13,000 ft. 
Range: Nepal and Sikkim. 

Ailurus fulgens styani Thomas, 1902 

1902. Ailurus fulgens styani Thomas, Ann. Mag. N.H. in: 251. Vangliupa, North- 

^\'estern Szechuan, China. 
?) 1874. Ailurus refulgens Milne-Edwards, Rech. Mamm. 380. 
R<uige: .Szechuan, ^'unu.m, Northern Burma. 

Genus AILUROPODA Milne-Edwards, 1870 

1870. Ailurnpoda Milne-Edwards, Ann. Sci. Nat. Zool. ij, art. 10: i. ihsus tnelano- 
leucus David. 

1870. Pandaretos Ger\'ais, Nouv. .\rch. Mus. H.N. Paris, G: 161. Ursus melannlcucus 


1 87 1. Ailuropus .Miliic-Fxlwards, in David, Nouv. Arch. Mus. H.N. Paris, 7, Bull. 92. 

I species: Ailiiro/ioda melanoleuea, page 243 


Ailuropoda melanoleuca David, i86g Giant Panda 

Approximate distribution of species: State of Szechuan, in China. 

Ailuropoda melanoleuca David, 1869 

i86g. Ursus melanoleucus David, Nouv. Arch. Mus. H.N. Paris, j. Bull. 13. Moupin, 
Szechuan, China. 


Genera: Aonyx, page 278 Aides, page 271 

Arcionyx, page 274 Mellivora, page 268 

Enhydra, page 279 Melogale, page 269 

Gulo, page 250 Mustela, page 251 

Luira, page 275 Poecilictis, page 267 

Martes, page 244 Vormela, page 266 

Pocock divided this large family into a bewildering number of subfamilies, which 
are reduced by Simpson (1945) so far as living Palaearctic and Indian genera are 
concerned, to four. These are the Mustelinae, the Melinae, and the Lutrinae, which 
are recognized by virtually all authors, and the Mellivorinae which does not seem 
strongly differentiated from Mustelinae. Simpson's arrangement is simpler than that 
of Pocock, and is followed here. Of the genera listed above, the characters of eight 
are dealt with by Pocock in his work on the mammals of British India (1941). Besides 
this he shows (p. 423) the distinguishing characters ofMeles compared with its nearest 
ally Arctonyx. Meles was also dealt with at some length by Miller, 1912, Cat. Mamm. 
Western Europe, 341, and in that work the characters of Gulo are given (p. 433). 
Miller referred Gulo to a distinct subfamily, whereas Pocock thought it was nearest 
the Martens. For the characters oi Enhydra see Pocock, 192 1, P.^-S- 803-837, "On 
the External Characters and Classification of the Mustelidae". In this work, Enhydra 
is given subfamily rank under the name "Lataxinae" (p. 830). On p. 835 of the same 
work, the characters of the African Ictonyx group of genera, to which Poecilictis belongs, 
are given; this group is also given subfamily rank. There has been an increasing 
tendency towards genus-splitting in this family during recent years. Even Simpson 
lists four more genera than are here admitted, and Pocock about the same number, 
but their extra genera do not agree. We retain here genera which are universally 
admitted, and prefer to regard the possible extra genera as subgenera. The only 
genus here retained which is not of \ery longstanding is Poecilictis, which used to be 
referred to the earlier-named Ictonyx Kaup, 1835, but which seems a distinct form with 
peculiarly enlarged bullae which distinguish it from Ictonyx and in all probability 
from all the other Palaearctic genera belonging to the Mustelinae as here understood. 
Pocock referred Martes to a special subfamily, which following Simpson is here 
referred to the Mustelinae. He also made a special subfamily for Helictis (which is 
antedated by Melogale and here referred to that genus), which Simpson placed in the 


Subfamily M u s t e 1 i n a e 

Genus MARTES Pincl, 1792 

1773. Maiiis Friscli, Xatur-systcm der \icrfuss. Thierc, 11 (see page 2). 

1792. Martci Pinel, Actcs Soc. d'H.N. Paris, /; 55. MarUs domestica Pinel --^ Muslela 

foina Erxleben. 
1821). ^ibt'Uina Kaup, Entw. Gesch. u. Nat. Syst. Europ. Thierw. /.■ 31, 34. Muslela 

zihiilina Linnaeus. 
181)",. Charronia Gray, P.Z.S. 108. Mustela flavigula Boddaert. Valid as a subgenus. 
11)28. Lamprogale Ognev, Mem. Soc. Amis. Sci. Nat. Mosc. No. 2, Zool. 26, 30. 

Proposed to replace Cliarronia on the grounds that it is preoccupied by an 

earHer name Ckaronia, for a genus of mollusc. 

This genus was formerly known as Alustcla by many authors, as, for instance, 
Blanford, 1891. What is now known as Mustela was called by older authors Putorius. 

6 species in the area covered by this list: 

Marks flavigula, page 249 
Marks foina, page 246 
Maries gwatkinsi, page 250 
Marks marks, page 245 
Marks inelampus, page 247 
Marks .-jbellina, page 248 

The species flavigula and gwatkinsi belong to the subgenus Charronia, which is fully 
compared with Maries by Pocock (1941, 319, 326). The characters of the two species 
are also dealt with by Pocock. Miller (1912) compares the characters of martes and 
foina, which apparently are not always very easily distinguishable. A translation in 
our possession of part of Ognev's work on the mammals of the U.S.S.R. contrasts 
M. -ihellina with M. marks and M. foina, and states that m zibelUna there are 15-16 
tail \crtebrac, whereas in the other two species there are 20 and more; also that in 
zibellina the bullae are differently shaped, and set closer together, as may be seen 
from Bobrinskii's figures of the three species' skulls (1944, 11 8). Bobrinskii states that 
zibellina has the tail usually less than half head and body length, the light patch on 
the throat is often absent, and the top of the head is usually lighter than the back, 
\\ hcrcas in martes and Joina the tail is usually more than half the head and body length, 
the- light spot on the throat is well developed, and the top of the head is the same 
colour as the back. In the London material, however, it must be noted that M. martes 
skins have the tail averaging only 49 per cent, of the head and body. There remains 
for discussion the Japanese species M. melampus. In the London material, this has the 
tail on average about 44-47 per cent, of head and body length {rcsemhlmii^ zibellina, 
therefore, in rather short tail) ; a white throat patch seems fairly constant, and, at least 
in winter, the head tends to be paler than the back, all characters reminiscent of 
Zibellina except the throat patch. But the bullae seem to be definitely of the martes- 
Joina type, and do not seem to resemble those of zibellina. The forelimbs are clearly 
contrasted blackish, more so than in our zibellina skins. Therefore the conclusion has 



been reached that melampus is an isolated and valid species, partly combining the 
characters of the other two groups. So far as colour is concerned it in no way resembles 
the subgenus Charronia as that is defined by Pocock. But it must be noted that not all 
our skins o^ melampus bear measurements. 

.Subgenus MARTES Pinel, 1792 

Martes martes Linnaeus, 1758 Pine Marten 

Approximate distribution of species: British Isles, Ireland included; Norway, 
Sweden, Denmark, France, Belgium, Holland, Germany, Switzerland, Italy, 
Northern Spain, Balearic Islands, Sardinia, Bohemia, Poland, to Russia, from White 
Sea to Caucasus, and eastwards into Western Siberia, roughly to lower Ob and lower 
Irtish Rivers. British Museum localities also include .Sumela (Asia Minor) and Astra- 
bad i' Persia). 

Martes martes martes Linnaeus, 1758 

1758. Mustela martes Linnaeus, Syst. Nat. loth ed. /.• 46. Upsala, Sweden. 
1816. Mustela sylvestris Oken, Lehrb. Nat. j, 2: 1029. Renaming oi martes. 
1820. Martes sjlvatica Nilsson, Skand. Fauna. Dagg. Djur. /.• 41. Renaming oi martes. 
1827. Martes vulgaris Griffith, Cuvier's Anim. Kingd. §: 123. Renaming oi martes. 
1865. Martes abietum Gray, P.Z.S. 104. 

Range: Europe, north of the Mediterranean; Russia, as far as the \Vhite Sea and a 
line from Kiev to Vitebsk. 

Martes martes latinorum Barrett-Hamilton, 1904 

1904. Mustela martes latinorum Barrett-Hamilton, Ann. Mag. N.H. i^: 389. Nurri 
Mountains, Sardinia. Range: Italy, Sardinia, Balearic Islands. 

Martes martes notialis Cavazza, 191 2 

1912. M{ustela) martes va.r. notialis Cavazza, Ann. Mus. Civ. Stor. Nat. Genova, 3A, 
5 (45): 181. South of Abruzzi, Southern Italy. 

Martes martes lorenzi Ognev, 1926 

1926. Martes martes lorenzi Ognev, Bull. Sci. Inst. Expl. Caucas. /.• 47. Storojevaia, 
Kuban district, Caucasus. 

Martes martes ruthena Ognev, 1926 

1926. Martes martes ruthena Ognev, Bull. Sci. Inst. Expl. Caucasus, /.■ 49, 56. Dmit- 

rovsk subdistrict, Moscow Govt., Russia. Range: Central regions of 

European Russia. 

Martes martes borealis "Kuznetzov, 1941," Bobrinskii, 1944 
1944. M{artes) m{artes) borealis Bobrinskii, Mamm. U.S.S.R. 121. Not of Radde, 
1862. Northern areas of European Russia, excluding Kola Peninsula. 



Martes martes uralensis "Kuznetzov, 1941," Bobrinskii, 1944 
1944. M{arles) m[arla) uralensis Bobrinskii, Mamm. U.S.S.R. 121. Whole area of the 
Ural Rane;e. 

We are unable to trace the original reference to the last two named forms. 

Martes foina Erxleben, 1777 Beech Marten, or Stone Marten 

Approximate distribution of species: Europe, including Spain, Italy, Bosnia, 
France, Belgium, Holland, Germany, Denmark, Switzerland, Crete, Poland, Fin- 
land (Ognev), Russia (Ukraine, Crimea, Caucasus, Transcaucasia (part) and Western 
Russia, according to Bobrinskii) ; Russian Turkestan (mountain areas), northwards to 
the Altai; Asia Minor, Persia, Afghanistan, Syria and Palestine; Baluchistan, 
Kashmir, Punjab; Chinese Turkestan, Mongolia, Manchuria (Bobrinskii), Tibet. 
Possibly parts Northern China. Perhaps to Sikkim. 

Martes foina eoina Erxleben, 1777 

1777. Miisida foina Erxleben, Syst. Regn. Anim. /.• 458. Germany. 

1792. Martes domestka Pinel, Actes Soc. H.N. Paris, /.• 55. France. 

1801. M I) s hi a foina alba Bechstein, Gemeinn. Nat. Deutschlands, 2nd ed. /.■ 759. 

Thuringia, Germany. 
1869. Mustela martes \a.Y. fagorum Fatio, Faune Vert. Suisse, /.• 318. 
Range: Europe, as above, except Southern Spain; probably eastwards into Russia. 

Martes (?) foina toufoeus Hodgson, 1842 

1842. Mustela? toufoeus Hodgson, J. Asiat. Soc. Bengal, //.• 281. ? Lhasa, Tibet. 
Despite Pocock's contention that this is allied to M. melampus, it looks much 
more like M. foina. Its range is adjacent to that of foina, very far from melampus. 
From notes left by him, Chaworth-Musters evidently intended to treat it as 
foina. See also Pocock (1941, 322, footnote). We cannot trace that the form 
"ka?isuensis" noted by him on this page was ever described. 

Martes foina intermedia Severtzov, 1873 

1873. Mustela intermedia Severtzov, Mem. Soc. Amis. Sci. Nat. Moscow, 8, 2: 61. 

1876, Ann. Mag. N.H. 18: 45. Basin of the Chu, Tallas, and Naryn, from 

4,000 to 9,000 ft.. Eastern Turkestan. 
1879. Martes leueolachnaea Blanford, Second Yarkand Miss. Mamm. 26. Varkand, 

Chinese Turkestan. 
1914. Martes foina altaica Satunin, Conspectus Mamm. Ross. /.• iii. Altai. 
1919. Martes toufoeus Wroughton, J. Bombay N.H. Soc. 26: 343. Not of Hodgson, 

Range: Russian and Chinese Turkestan, Tianshan, Afghanistan, Baluchistan, 
Western Persia, Kashmir. 

Martes foina mediterrane.\ Barrett-Hamilton, 1898 

1898. Mustela mediterranea Barrett-Hamilton, .A.nn. Mag. N.H. /.• 442'. Sierra de 
Jerez, Cadiz, .Spain. 



Martes foina syriaca Nehring, 1902 

1902. Mustela foina syriaca Nehring, S.B. Ges. Nat. Fr. Berlin, 145. Wadi Syr (which 
runs into Wadi Kefren, a tributary of lower Jordan), Syria. 

Martes foina bunites Bate, 1906 

1906. Mustela foina bunites Bate, P.Z.S. igo§, 2: 318. Kontopalo, Kania, Crete. 

Martes foina nehringi Satunin, 1906 

1906. Mustela foina nehringi Satunin, Mitt. Kauk. Mus. Tiflis, 2: 120, 292. Tiflis, 

Martes foina bosniaca Brass, i g 1 1 

191 1. Martes foina bosniaca Brass, Aus der Reiche der Pelze, 468 (spelt "bosnia'' in 
index, p. xiii). Bosnia, Yugoslavia. 

Martes foina milleri Festa, 1914 

1914. Martes foina milleri Festa, Boll. Mus. Zool. Anat. Comp. Torino, 2g, 686: 7. 
Aghios Isidoros, Island of Rhodes, Eastern Mediterranean. 

Martes foina rosanowi Martino, 19 17 

1 91 7. Martes rosanowi Martino, Bull. Soc. Nat. Crimee, y: i. (Reprint only seen.) 
North-western slope of Chatyr dag Mountains, Crimea, Southern Russia. 

Martes foina kozlovi Ognev, 1931 

1 93 1. Martes foina kozlovi Ognev, Mamm. E. Europe, N. Asia, 2: 631. Kam (valley 
of River Mekong), Eastern Tibet. 

Martes melampus Wagner, 1841 Japanese Marten 

Approximate distribution of species: Japan (Hondo, Shikoku, Kiushiu, Tsushima) 
and Korea. 

Martes melampus melampus Wagner, 1841 

1841. Mustela melampus Wagner, Schreb. Saugeth. Suppl. 2: 229. Japan. 
1865. Martes japonica Gray, P.Z.S. 104. Japan. 
1865. Martes melanopus Gray, P.Z.S. 105. 

1905. Mustela melampus bedfordi Thomas, Abstr. P.Z.S. 10, P.Z.S. 2: 183. Washika- 
guchi, Nara district, east of Osaka, Southern Hondo, Japan. 

Martes melampus tsuensis Thomas, 1897 

1897. Mustela melampus tsuensis Thomas, Ann. Mag. N.H. ig: 161. Kamoze, 
Tsushima Islands, Japan. 

Martes melampus coreensis Kuroda & Mori, 1923 

1923. Martes melampus coreensis Kuroda & Mori, J. Mamm. 4: 27. Tenan, Southern 
Chusei district, Korea. 



Martes zibellina Linnaeus, i 758 Sable 

Approximate distribution ofspccies: from the Pcch<ira River and Ural Mountains, 
eastwards intermittently throu£jh Siberia to Kamtchatka, Sakhalin and the Ussuri 
region, south to the Altai Mountains, mirth to the Arctic Circle, and somewhat 
beyond in Middle Siberia. Manchuria, Mongolia and Japan. (Now only surviving 
in indi\idual isolated areas, Bobrinskii.) 

Martes zibeli.ina zjbejaasa Linnaeus, 1758 

1758. Miisli'la zil'elli'id Linnaeus, Syst. Nat. loth cd. /.' 46. Surroimdings of Tobolsk, 

Tomsk Govt., Siberia (Ognev). 
185-,. Miistcla Z'beHina \'ar. asiatica Brandt, Mem. Phys. Nat. Acad. Sci. St. Petcrsb. 

;.- 6, pi. I. 
185",. Mustela .zibellina \ar. aiha Brandt, loc. at. j: 14, pi. 2, fig. 5. 
1855. Miislela zil'fllina var. Jiisco-Jiavescens Brandt, loc. cit. pi. 2, fig. 6. 
1855. Mustela zibellinn var. ochracea or ferruginea Brandt, loe. cit. \i\. 3, fig. 8. 
1855. Mustela zibellina var. maculata Brandt, loc. cit. pi. 3, fig. g. 
1855. Mustela .zibellina \'ar. rupcstris Brandt, loc. cit. pi. 2. 
1855. Mustela zibellina var. svlvestris Brandt, loe. cit. pi. 2. 
Range: Pechora basin. Northern Urals, Ob plain. 


1844. Mustela brachyura Tcmminck, Sicbolds Faun. Japon. Mamm. 33. Japan. 
(Veso — Hokkaido and the Kurilos.) 


IQ18. Mustela zibellina subsp. kamtshadalica Birula, C.R. Mus. Zool. Acad. Sci. 

Petrogr. 82. [j\.V. Reference according to Ognev.) Kamtchatka. 
(?) 1922. Mustela zibellina var. kamtschatica Dybowski, Arch. Tow. Nauk. Iavow, i: 

349, nom. nud. 


1922. Mustela zibellina princeps Birula, Ann. Mus. Zool. Acad. Sci. St. Petersb. 2::: 8. 

Bargusin Mountains, Transbaikalia. 
?) 1922. Mustela zibellina var. baiealcnsis Dybowski, Arch. Tow. Nauk. Lwovv, i: 
349, nom. nud. 


1925. Martes zibellina veniseensis Ognev, J. NLimm. 6: 277. Forest on plain along 
Ycnesei River, Krasnoiarsk district. Eastern Siberia. Range: Taiga between 
the .Angarra and the Sayan foothills fBobrinskii). 

Martes zibellina sajanensis Ognev, 1925 

1925. Martes zibellina sajanensis Ognev, J. Mamm. G: 278. Orsyba River, northern 
part of Sayan Mountains, Midfllc Siberia. 

M.-\RTEs zibellina s.\halinensis Ognev, 1925 

192',. Maries zibellina sahalinensis Ognev, J. Mamm. (j: ^ji). \\'ederniko\o, Sakhalin 



Martes zibellina hamgyenensis Kishida, 1927 

1927. Martes zibellina coreensis Kishida, Choju Chosahokoku, 4: 130. Korea. Not of 

Kuroda & Mori, 1923. [N.V.) 
1927. Martes zibellina hamgyenensis Kishida, Dobuts Zasshi. jp; 509 {N.V.) 
1931. Martes zibellina hangiengensis Kishida & Mori, op. cit. 43: 380, nom. mid. (N.V.) 

These references are from Kuroda. 

Martes zibellina tungusensis "Kuznetzov, 1941," Bobrinskii, 1944 
1944. M[artes) z(ibellina) tungusensis Bobrinskii, Mamm. U.S.S.R. 120. Basins of the 
Nizhnaya and Podkamennaya Tungusha (Middle Siberia). 

Martes zibellina arsenjevi "Kuznctzov, 1941," Bobrinskii, 1944 
1944. M{artes) z{ibellina) arsenjevi Bobrinskii, Mamm. U.S.S.R. 120. Ussuri basin, 
Eastern Siberia. 

Martes zibellina schantarica "Kuznetzov, 1941," Bobrinskii, 1944 

1944. Miartes) z{ibellina) schantaricus Bobrinskii, Mamm. U.S.S.R. 120. Shantar 

Islands, Lower Amur, Eastern Siberia. 
(?) 1922. Mustela zibellina var. amurensis Dybowski, Arch. Tow. Nauk. Lwow, /.• 349, 

nom. mid. 

We are unable to trace the original reference to the last three listed races, which 
are without description in Bobrinskii, 1944. 

Subgenus CHARRONIA Gray, 1865 [Lamprogale Ognev, 1928) 

Martes flavigula Boddaert, 1785 Yellow-throated Marten 

Appro.ximate distribution of species: Amur and Ussuri regions of Eastern Siberia; 
Korea, Manchuria, throughout the principal states of China (Chihli, perhaps, 
excepted), Tibet, Formosa; Burma, Assam, thence westwards to Kashmir and North- 
\Vest Frontier; Indo-China, Siam, Malay States, Sumatra, Java and Borneo. 

Martes flavigula flavigula Boddaert, 1785 

1785. Mustela Jlavigula Boddaert, Elcnch. Anim. 88. Locality ujiknown, traditionally 

fixed as Nepal (Pocock). 
1792. Mustela melina Kerr, Anim. Kingd. 183. Locality unknown. 
1800. Viverra quadricolor Shaw, Gen. Zool. Mamm. /, 2: 429. Locality unknown. 
1800. Mustela leucotis Bechstein, Uebers. vierf. Thiere, 2: 375. Locality unknown. 
1828. Mustela hardwickei Horsfield, Zool. J. 4: 239, pi. 8. Nepal. 
1842. Galidictis chrysogaster H. Smith, Jardine's Nat. Lib. jj, Mamm. i: 167. Mus- 

soorie, Kumaon, Northern India. 
1901. Mustela flavigula typiea Bonhote, Ann. Mag. N.H. y: 343. 
1901. Mustela flavigula kiiatunensis Bonhote, Ann. Mag. N.H. j: 348. Kuatun, North- 

Western Fukien, Southern China. 
1910. Mustela flavigula szetchuensis Hilzheimer, Zool. Anz. jjj.- 310. Sungpanting, 

Szechuan, China. 




1922. Charronia melli Matschie, in Mcll, Arch. Nat. 88, sect. A, 10: 17, 34. Kwantung, 

Southern China. 
1930. Charronia yuenshanensis Shih, Bull. Dept. Biol. Sun Yatsen Univ. Canton, No. 9, 

3. Yuen Shan, Wuchanghsien, Hunan, China. 
Range: Kashmir to Tibet and Southern China, north to Shcnsi, Kansu. 

Martes flavigul.'^ aterrim.^ Pallas, 181 1 

181 I. Viverra aterrima Pallas, Zoographia, /.- 81. Between the Uth and Amur Rivers, 
Eastern Siberia. 

1862. Mustela (Martes)-flavigula var. borealis Raddc, Reise Ost. Sib. /.• 19, 24. Moun- 
tains of Bureinsk, Siberia. 

1922. Charronia flavi gill a koreana Mori, Ann. Mag. N.H. 10: 610. Korio. near Seoul, 

M.-\RTES FL.'^viGULA CHRYSospiLA Swinhoe, 1 866 

1866. Marlfs chrysospila Swinhoe, Ann. Mag. N.H. 18: 286. Mountain forests of 

Central Formosa. 
1870. Martes flavigida xan/hospila Swinhoe, P.Z.S. 623. Forests of Central Mountains 

of Formosa. 

Martes fl.'^vigula pemnsul.^ris Bonhote, 1901 

iqoi. Mustela flavigula pemmiilaris Bonhote, Ann. Mag. N.H. y: 346. Bankasun, 
Tenasserim. Range: to Malay Peninsula. 

Martes flavigula lxdochinEiNSIs Kloss, 1916 

1 916. Martes flavigula indochinensis Kloss, P.Z.S. 35. Klong Menao, South-Eastern 
Siam. Range: Northern Tenasserim, Siam, Annam. 

Martes gwatkinsi Horsfield, 1851 South Indian Yellow-throated Marten 

Approximate distribution of species: Nilgiri Hills, Coorg and Travancore, 
Southern India. 

Martes gwatkinsi Horsfield, 1851 

1851. Martes gwatkinsii Horsfield, Cat. Mamm. E. Ind. Co. 99. Madras, Indi;i. 

Genus GULO Storr, 1780 

1775. Culo Frisch, Natur-system der vierfuss. Thiere, 17 see page 2). 
1780. Giilo Storr, Prodr. .\ieth. Mamm. 34. Tab. \. Mustela gulo Linnaeus. 

I species: Gulo gulo, page 250 

Gulo gulo Linnaeus, 1758 Glutton, or Wolverine 

Approximate distribution of species; Norway and Sweden; "right across the taiga 

and forest-tundra zone of Eastern Europe, Asia and North America. In the summer 

it invades the tundra, as far as the sea coast. In Eastern Eumpe and Western Siberia 


it extends roughly as far south as the latitude of Sverdlovsk, but occurs in an isolated 
area south-west of Kiev. In the more eastern parts of Asia it extends south to the 
Altai, Tuva Republic, Mongolia and Northern Manchuria, inclusive. It does not 
occur in the Transbaikal steppes. In the south of the Ussuri region it is rare. It occurs 
in Sakhalin and the Shantar Islands" (Bobrinskii). 

GuLO GULO GULO Linnaeus, 1758 

1758. Mustela gulo Linnaeus, Syst. Nat. loth ed. /.■ 45. Lapland. 

1780. Gulo sibirica Pallas, Spic. Zool. 2, 14: 35, Tab. 2. 

1792. Ursus gulo albus Kerr, Anim. Kingd. Syst. Cat. No. 381, igo. Kamtchatka. 

1816. Gulo vulgaris Oken, Lehrb. Nat. 3, 2: 1004. Renaming oi gulo. 

1820. Gulo borealis Nilsson, Skand. Faun. Dagg. Djur. /.• 95. Renaming o{ gulo. 

1820. Gulo arcticus Desmarest, Mammalogie, 174. Renaming oi gulo. 

1829. Gulo arctos Kaup, Entw. Gesch. Nat. Syst. Europ. Thierw. /.• 68. Renaming of 

1 9 10. Gulo luscus Trouessart, Faune Mamm. d'Europ. 71. Not of Linnaeus,. 1766. 
1 918. Gulo biedermanni Matschie, S.B. Ges. Nat. Fr. Berlin, 147. Mountains south of 

Lake Teletzkoie, Siberian Altai. 
1918. Gulo ivachei Matschie, op. cit. 147. North of Beluha Mountains, in upper reaches 

of River Katun, Siberian Altai. 
1922. Gulo kamtschaticus Dybowsky, Arch. Tow. Nauk. Lwow, /.• 349, nam. nud. 


Genus MUSTELA Linnaeus, 1758 

1758. Mustela Linnaeus, Syst. Nat. loth ed. /.• 45. Mustela erminea Linnaeus. 

1775. Putorius Frisch, Natur-system der vierfuss. Thiere, 11 (see page 2). 

1817. Putorius Cuvier, Regne Anim. /.• 147. Mustela putorius Linnaeus. \'alid as a 

1829. Arctogale Kaup, Entw. Gesch. Nat. Syst. Europ. Thierw. /.• 30. Mustela erminea 

1829. Ictis Kaup, Entw. Gesch. Nat. Syst. Europ. Thierw. /.• 35, 40, 41. Mustela 

vulgaris Erxleben = Mustela nivalis Linnaeus. Not of Schinz, 1824-1828. 

1840. Foetorius Keyserling & Blasius, Wirbelth. Europ. 68. Mustela putorius Linnaeu;. 

1841. Gale Wagner, Schreb. Saugeth. Suppl. 2: 234. Mustela vulgaris Erxleben = 

Mustela nivalis Linnaeus. 
1841. Lutreola Wagner, Schreb. Saugeth. Suppl. 2: 239. Viverra lutreola Linnaeus. 
1865. Gymnopus Gray, P.Z.S. 118. Mustela leucocephalus Gray = Alustela nudipes 

Desmarest. Not of Brookes, 1828. 
1871. Mustelina Bogdanov, Proc. Imp. Univ. Kazan, /.• 167. Mustela lutreola Linnaeus. 
1 87 1. Hydromustela Bogdanov, Proc. Imp. Univ. Kazan, /.• 167. Mustela lutreola 

1899. Eumustela Acloque, Faune de France, Mamm. 62. Based on vulgaris and erminea. 
igii. Kolonokus Satunin, Mitt. Kauk. Mus. j.- 264. Mustela sibirica Pallas. 
1 92 1. Plesiogale Pocock, P.Z.S. 805. Mustela nudipes Cuvier. Not of Pomel, 1853. 
1947. Pocockictis Kretzoi, Ann. H.N. Mus. Hung, ^o.- 285. To replace Plesiogale 

Pocock. Mustela nudipes C\i\'\ev. 


8 species in the area covered by this hst: 

Mustela altaica, page 259 Muslela nivalis, page 256 

Mustela erminca, page 253 Mustda putorius, page 264 

Muslela kathiali, page 259 Mustela sibirica, page 260 

Muslela hihrola, page 262 Muslela strigidorsa, page 264 

Miller, H)I2, Cat. Mamiii. Western Europe, 382, divided Mustela into three sub- 
genera, Mustela. Lutreola and Putorius, and gave characters for the three groups. These 
subgenera are recognized by Bobrinskii, 1944, Mammals of the U.S.S.R., and the 
characters given by Miller are more or less confirmed. Pocock, on the other hand, 
gave Putorius generic'rank, and appeared to ignore Lutreola. 

Russian authors recognize two species of the subgenus Putorius, which are said to 
occur together in some places: M. putorius, characterized by having the tail nearly all 
black, the underparts blackish, black predominating on upper side of body, and the 
skull with hardly any postorbital constriction; and M. eversnianni , with only the 
terminal half of the tail black, the underparts light-coloured, the upper parts with 
vcllowish straw-colour predominating, and the skull with a marked postorbital con- 
striction. The Ferret, ^[. p. furo of Linnaeus, antedates eversmanni, but in external 
appearance seems to agree more with eversmanni than putorius. Pocock thought it was 
a semi-domesticated descendant of putorius, and stated that its skull was like that of 
putorius, not eversmanni. He thoroughly reviewed the group, 1936, P.^.S. 691, and 
came to the conclusion that all members of the subgenus Putorius are one species. 
According to Bohrinskii, putorius and eversmanni inhabit diflerent types of country, the 
latter "avoids both woodland areas densely grown with bushes, and human settle- 
ments", unlike putorius. On account of intermediate characters within the subgenus, 
we prefer tentatively to fallow Pocock and list all Polecats in one species only, 
M. putorius. 

An attempt to correlate the work of Miller, Bobrinskii, G. Allen and Pocock with 
regard to the species of the subgenus Mustela, and to include outlying forms not dealt 
with by those authors, as, for instance, from Japan, South-^Vestern Asia (where the 
subgenus is rare) and North Africa leads to the f lUowing provisional results. M. 
stngidorsa differs from all others by its narrow, pale middorsal stripe. The soles of its 
feet are naked. The two species lutreola and sibirica stand somewhat apart from the 
remainder in that the underparts are only very little paler, if at all, than the upper 
parts. [M. sibirica can ha\e a white chin.) They differ from each other cranially, as 
noted by Miller (191 2) [Lutreola, subgenus, for .U. lutreola, while sibirica appears to 
agree with subgenus Mustela) ; and as figured by Bobrinskii ( 1 944, 1 24) . In the remain- 
ing species, the underparts are normally clearly lighter coloured than the upperparts, 
except of course in the winter coat of those forms which turn completely white. 
.U. erminea is characterized by its very sharply contrasted black tailtip, which is re- 
tained even in the wholly white winter coat when present. We prefer to regard the 
oullving .\/. hibernica from Ireland as a race of erminea. As discussed below, it appears 
that ermineu oc( urs in Algeria. There remain the nivalis group and the altaica group. 
In these, the IjKk k tailtip is usually absent or is represented by a few dark hairs only 
at the end ol the tail. In the Eastern Asiatic .\/. altaica and .\L kathiah the tail ap]jears 


not specially shortened, and so far as we can discover is nearly always at least 
100 mm. in length. We do not think that G. Allen was correct in making kathiah a 
subspecies oi altaica, as the two seem to occur in the same general neighbourhood in 
Himalayan India. Pocock has given characters to separate the two species, and we 
retain kathiah. In M. nivalis the colour is not Very different from the altaica type, but 
the tail is normally very shortened, being less than loo mm. in length so far as is 
ascertainable, except in North Africa. A broad view is here taken of the species M. 
nivalis. Some authors prefer to regard some of the eastern races as subspecies of the 
North American AI. rixosa Bangs (Putorius rixosus Bangs, 1896, Proc. Biol. Soc. Washing- 
ton, 10: 21, Saskatchewan, Canada), but Bobrinskii refers all the Russian and 
Siberian weasels to M. nivalis. The possibility that there is a larger and a smaller 
species of the nivalis group cannot however be finally dismissed. The names suhpalmala 
(1832, Egypt), numidica {1855, Morocco) and algirica (1895, Algeria) are available for 
the North African members of the subgenus Mustela. The first is obviously a large 
member of the nivalis group, in that there is no suspicion of a black tailtip. The 
second has a short black tailtip, according to Cabrera, but as figured by him looks 
more like subpalmata, the dark tip being poorly contrasted, and a specimen in the 
British Museum from IVIorocco seems to have no black tip. The form algirica was 
described by Thomas as a race of A/, erminea, and certainly seems to be, on account of 
the black tailtip. Its feet, also, are whiter than our other North African skins. 
Cabrera, and following him G. Allen, placed it in synonymy oi numidica. The question 
cannot be settled without more specimens, but if it is a synonym, then numidica repre- 
sents erminea, and if not, then both nivalis and erminea occur in North-West Africa. 
Tentatively, the latter conclusion has been adopfed. The large Egyptian weasel 
[subpalmata) can have the tail over 100 mm. in length, though it seems always under 
half the head and body length in our specimens, which is not normal in M. altaica and 
M. kathiah so far as measurements of these are available. 

Cranial characters used by Bobrinskii to separate M. altaica and sihirica from M. 
nivalis and erminea are not constant in the British Museum material when specimens 
from outside the U.S.S.R. are considered. 

Of other outlying forms, M. itatsi, Japan, often given specific rank, does not seem 
certainly separable from M. sibirica. The form stoliczkana (Yarkand) and the small 
form russelliana fSzechuan) seem to represent the nivalis group, and the recently 
described tonkinensis (Indo-China) may also be a largish southern member of the same 
group; it is not a representative oi kathiah, since the latter occurs in the same area. 

The only other species in Asia is M. nudipes Cuvier, 182 1, from Sumatra, Malava 
and Borneo; on this, see Pocock, 1941, Fauna Brit. India. 2: 379. 

Subgenus MUSTELA Linnaeus, 1 758 
Mustela erminea group 

Mustela erminea Linnaeus, 1758 Stoat ^Ermine) 

Approximate distribution of species: Europe, from Arctic south to Pyrenees -and 
Alps (including British Isles, west to Ireland, Sweden, Norway, France, Belgium, 



Holland, Denmark, Switzerland, Germany, Poland, Czechoslovakia) ; Russian range 
Efiven by Bobrinskii as "whole of Eastern Europe except Novaya Zemlya and Crimea. 
The Northern Caucasus, where it is very rare. Does not occur in Transcaucasia. 
\\'hole of Siberia to the Shantar Islands and Sakhalin. Kotelnuii Island (Novo- 
sibirskie group). Kazakstan (except for the extreme south), Kirghizia and 
Tadzhikistan". Mongolian .\ltai, Kashgaria, Japan; Afghanistan; Kashmir; Algeria. 
.\lso in North America. 

MiiSTEL.\ ERMINEA ERMi.\E.\ Linnacus, 1758 

1758. Mustela erminca Linnaeus, Syst. Nat. loth cd. /.• 46. Sweden. 

17Q2. Mustela t-rminea hvherna Kerr, Anim. Kingd. 181. 

1816. Mustela herminea Oken, Lehrb. Nat. 5, 2: 1026. Renaming of erminea. 

1827. Mustela erminea maculata Billberg, .Synops. Faun. Scandinaviae, 8. Scandinavia. 

Range: Norway, Sweden, Kola Peninsula in Northern Russia. 

Mustela erminea aestiv,\ Kerr, 1792 

1792. Mustela erminea aestiva Kerr, Anim. Kingd. 181. Germany. 

1820. Mustela erminea major Nilsson, Skand. Faun. Dagg. Djur. /.• 34. Carlskrone, 

Blekinge, Sweden. 
(?) 1920. Putorius ermineus giganteus Burg, Der W'cidmann, 48, 388. (M.V.) 
(?) 1920. Putorius ermineus alpestris Burg, he. cit. [N.W) 

P.ange: Continental Europe, from Southern Sweden south to Alps and Pyrenees, 
eastwards through Russia to Kazakstan. 

Mustela erminea hibernic.\ Thomas & Barrett-Hamilton, 1895 

1895. Putorius hihernicus Thomas & Barrett-Hamilton, Ann. Mag. N.H. /j.- 374. 
Enniskillen, Co. Fermanagh, Ireland. Range includes the Isle of Man. 

MusTEL.'V erminea algiric.\ Thomas, 1895 

1895. Putorius ermineus algirieus Thomas, Ann. Mag. N.H. i§: 451. Near Algiers, 

Mustela erminea ferghanae Thomas, 1895 

1895. Putorius ermineus ferghanae Thomas, Ann. Mag. N.H. i§: 452. Mt. Kara Karyk, 

Ferghana, Eastern Russian Turkestan. 
1908. Mustela ivhiteheadi Wroughton, J. Bombay N.H. Soc. 18: 882. Kaghan Valley, 

Hazara, Northern India. 
Range: Eastern Russian Turkestan, southwards to Kashmir; also, according to 
Ognev, Kashgar and Afghanistan. 

Mustela .vrctica Merriam, 1896 

1896. Putorius arctuus .Merriam, North Amer. Fauna, //.• 15. Point Barrow, Alaska. 
(?) 1922. Putorius eiminea \\\r, kamtschatiea Dybowski, Arch. Tow. Nauk. Lwow, /.• 

341), nom. iiud. 
(?J 1944. Mu\tela etmmea digna Hall, Proc. C^alif. .Acad. Sci. I'j; 559. Kamtchatka. 



MusTELA ERMINEA STABiLls Barrctt-Hamilton, 1904 

1904. Putorius ermineus stabilis BaTTett-Hamihon, Ann. Mag. N.H. 13: 394. Blandford, 
Dorset, England. Range: mainland of Great Britain. 


1907. Putorius erminea ricinae Miller, Ann. Mag. N.H. 20: 395. Islay House, Island of 
Islay, Hebrides. Range also includes Island of Jura, Hebrides. 


191 2. P{utorius) ermineus var. minimus Cavazza, Ann. Mus. Civ. Stor. Nat. Genova, 
3A, 5 (45): 194. Monte Rosa, Switzerland. 


191 2. Mustela lymani Hollister, Smiths. Misc. Coll. 60, 14: 5. Tapucha, Altai Moun- 

tains, Siberia. 

Mustela erminea Nippon Cabrera, 1913 

1913. Mustela nippon Cabrera, Bol. Soc. Esp. /j.- 392. Sinano, Hondo, Japan. 

Mustela erminea tobolica Ognev, 1923 

1923. Arctogale erminea tobolica Ognev, Biol. Mitt. TimiriazefF, /.• 112. Tara, Tobolsk 
Govt., Western Siberia. 

Mustela erminea transbaikalica Ognev, 1928 

1928. Mustela erminea transbaikalica Ognev, Mem. Soc. Amis. Sci. Nat. Moscou, Sect. 

Zool. 2: 14, 29. Sosnovka, Bargusin forest, east shore of Lake Baikal, 


Mustela erminea orientalis Ognev, 1928 

1928. Mustela erminea orientalis Ognev, Mem. Soc. Amis. Sci. Nat. Moscou, Sect. 
Zool. 2: 15, 29. Village Pochodskoie, Kolyma River, North-Eastern Siberia. 

1914. Mustela kanei G. Allen, Proc. New Engl. Zool. Club, 5.' 58. Nijni Kolymsk, 

Eastern Siberia. Not of Baird, 1857. Recorded from Sakhalin, Kuriles 
and Hokkaido. But see also Hall, 1944, Proc. Calif Acad. Sci. 23: 555. 

Mustela erminea mongolica Ognev, 1928 

1928. Mustela erminea mongolica Ognev, Mem. Sect. Zool. Amis. Sci. Nat. Moscou, 2: 

18, 29. Dundu-Saichan, Mongolian Altai. 

Mustela erminea baturini Ognev, 1929 

1929. Mustela erminea baturini Ognev, Bull. Pacif Sta. Vladivostock, 2, 5: 9, 40. 

Bolshoi Shantar Island, Eastern Siberia. 

Mustela erminea ognevi Jurgenson, 1932 

1932. Mustela erminea o^««)i Jurgenson, Zool. Anz. g8: 1 1 . Delta of River Tas, extreme 
north of Central Siberia. 

Mustela erminea shnitnikovi Ognev, 1935 

1935. Mustela erminea shnitnikovi Ognev, Mamm. U.S.S.R. j.' 37. Kopal district, 
Semirechyia, Eastern Russian Turkestan. 



1036. Aliistila mninca karaoinensis ]uTs,civm, Hull. Soc. Nat. Moscciu, Sec. Biol. 4^: 
•2^0, -J.!;. Karaoinski Island, otT iKirth-cast coast of Kamtchatka. 


103B. Miislilii iiminca //(///mor; Jurscnson, Trav. Res. Etat. Altai, /.• 124. Source of 
the Khataiit;a, Tuiukhansk district (Northern Yenesei), Siberia. 


11)31. Miiilihi tiniinai hirulai Martino, Ann. Mus. Zool. Acad. Lcnin<;rad, ^i: J08. 
Aktiuhinsk, Kir<j;hi/ Steppes, Russian Asia. Not of Oc;nc\', 1928. 

Incertae ii-dis 

Putorius ermiiK 11 wir. w7)/«('(; Dybowski, 1922, Arch. Tow. Nauk. Lwovv, /.• 349, ito/n. 
mid. Not of Pallas, 1773. 

MnUila nivalis S>'oup 

Mustela nivalis l.iiin.ieus, 1766 Weasel. Type of G'a/c Wagner, 1841, 

if further subgeneric di\ision 
of the genus is required. 
.Appro.ximalc di^triliution of species : Europe, including Britain, France, Belgium, 
Holland, Denmark, Norway, Sweden, Germany, Poland, Switzerland, Portugal. 
.Spain, Italy, Sicily, Yugoslavia, Rumania; also Sardinia, Malta and Crete. In the 
U.S.S.R., the whole Union, according to Bobrinskii, although its presence has not 
been established in the north of the Taimyr Peninsula, and the interior of Kizil-Kum 
and Kara-Kum dcscits, and it does not occur in the islands of the Arctic Ocean. Asia 
Minor; Afghanistan; Mongolia, Koiea, Japan; Chinese Turkestan; Szechuan, in 
China; Egypt, Algeria, Morocco; ? Indo-China. Perhaps also in North America. 

MusTEi-.x xiv.\i,rs MVAijs Linnaeus, 1766 

1766. Mu\ltla nivalis Linnaeus, .Syst. Nat. 12th ix\. 1: bi). Pro\ince of Wsterbotten, 

1777. Miiilila vulgaris Erxleben, Syst. Rcgn. Anim. /.• 471. "Temperate Europe." 

Listed as a \alid race by Ognev, 1935, Mamm. L..S..S.R. 3'.' f)8, lor Southern 

Rirssian li)calitics, but considered a synonym by Miller, 1912. 
iBi I. Mii\h'la ,»(//(' Pallas, Zoogr. Ro.sso-Asiat. 94. Renaming ol vulgaris. 
1820. Muililii minor Nilsson, .Skand. Fauna, /.• 35. Renaming of nivalis. 
1853. Putorius miiiulus Pcjmel, C'at. Meth. et Descr. \'ert. Foss. Loire, 51. Near Paris, 

1869. Foelorius pusillus Fatio, Faune \'ert. Suisse, /.• 332. Not of De Kay, 1842. 
1900. Putonu\ nivalis Ivpicus Barrett-Hamilton, Ann. Mag. N.H. ^: 42. 
1908. Putorius nivalis \ar. monticola Cavazza, Richerche sui ^'Putorius nivalis" e sui 

"Putorius irrniiii'us" D'ltalia, 37 iM.V., see Miller, 1912, 412). High valle)s of 

the Alps. 
Range: Europe, fnjm ,\rctic coast to Alps and Pyrenees, and from Britain eastwards 
into Russia. 




1800. Mustela boccamela Bechstein, Pennant, Ubers. vierf. Thiere, 2: 395. Sardinia. 
(?) 1868. {Mustela vulgaris) var. fulva Mina Palumbo, Ann. Agric. Sicil. 12: 53. 

(JV.F.) Probably Le Madonie, Sicily. See Miller, 1913, Proc. Biol. Soc. 

Washington, 26: 80. Not of Kerr, 1792. 
(?) 1868. {Mustela vulgaris) var. albipes Mina Palumbo, loc. cit. 54. 
1869. Mustela vulgaris var. meridionalis Costa, Ann. Mus. Zool. della R. Univ. di 

Napoli, 40. Southern Italy. 
1900. Putorius nivalis italicus Barrett-Hamilton, Ann. Mag. N.H. 5.- 45. Grezzana, 

highlands of Verona, Italy. 

1900. Putorius nivalis siculus Barrett-Hamilton, Ann. Mag. N.H. j.- 46. Marsala, 


1 90 1. Mustela ilctis) dombrowskii Matschie, S.B. Ges. Nat. Fr. Berlin, 231. Siulnita, 

Rumania. Ognev thinks that this is a synonym of vulgaris, which is listed 

above under AI. nivalis nivalis. 
1905. Foetorius pusillus major Fatio, Arch. Sci. Phvs. Nat. Geneve, ig, 4: 512. 

Poschiavo, Grisons, Switzerland. Not of Nilsson, 1820. 
(?) 1920. Putorius boccamela alpinus Burg, Der Weidmann, 51, 409. {N.V.) 

Range: Italy, south coast of France, Sicily, Malta, Sardinia, Switzerland, Rumania. 

Mustela nivalis subpalmata Hemprich & Ehrenberg, 1833 

1833. Mustela subpalmata Hemprich & Ehrenberg, Symb. Phys. Mamm. 3, 2, k verso. 
In houses, Cairo and Alexandria, Eg>'pt. 

Mustela nivalis numidica Pucheran, 1855 

1855. Putorius numidicus Pucheran, Rev. Mag. Zool. j: 393. Tangier, Morocco. 

1865. Mustela erminea var. (i), africana Gray, P.Z.S. 1 1 1. Algiers, Algeria. Not africana 
Desmarest, 1818, Nouv. Diet. H.N. ig: 376, which is shown by Cabrera, 
1 914, to have been based on a South American species. 

1904. Putorius nivalis atlas Barrett-Hamilton, Ann. Mag. N.H. 13: 323. Atlas Moun- 
tains, Morocco. 

(?) 1908. Putorius nivalis var. corsicanus Cavazza, Ricerche sui "Putorius nivalis" e sui 
'"Putorius ermineus" d'ltalia, 37. Corsica. {N.V. See Miller, igi2, 412.) 

Range: Morocco, Algeria, Malta, Azores, ? Corsica. Miller suggests it was intro- 
duced in Malta and the Azores. Both Miller and G. Allen (1939) give this large 
form specific status. 

Mustela nivalis stoliczkan.\ Blanford, 1877 

1877. Mustela stoliczkana'QXa.nioTA,]. As\3.t. Soc. Bengal, 46, 2: 260. Yarkand, Chinese 
Turkestan. Ognev also quoted it from Djarkent (Eastern Russian Turkestan) 
and the Gobi, and it occurs Afghanistan (B.M.). 

Mustela nivalis nikolskii Smirnov, 1899 

1899. Foetorius vulgaris var. nikolskii Smirnov, Poslonjivotn. Krymea, 59 (appendix to 
68, Zap. Imp. Akad. Nauk). (.^V.r.) Near Simferopol, Crimea, Southern 


I'ALAEARtrriC; and INDIAN MAMMALS 1758-1946 

MusTELA NIVALIS iBERiCA Barrctt-Hamiltoii, 1900 

igoo. Puloriiis nivalis ibericus Barrett-Hamilton, Ann. Mag. N.H. 5.- 45. Seville, Spain. 
Range inrludcs Portugal and Balearic Isles. 

MusTELA NIVALIS PALLIDA Barrett-Hamilton, 1900 

1900. Puloriiis nivalis pallidiis Barrett-Hamilton, Ann. Mag. N.H. 5.- 48. Kokand, 
Ferghana, Eastern Russian Turkestan. 

MusTELA NIVALIS c.'^uc.xsiCA Barrett-Haniiltou, 1900 

1900. Piitoriiis nivalis caiicasiciis Barrett-Hamilton, Ann. Mag. N.H. §: 48. Hotshal 
Mountains, iQ,aot) ft., Caucasus. 


1903. Putorius [Arclogale] firgmaeusj. Allen, Bull. Amer. Mus. N.H iij: 176. Gicliiga. 

west coast of Okhotsk Sea, Eastern Siberia. 
(?) 1922. Ictis nivalis var. kamtschatica Dybowski, Arch. Tow. Nauk. Lwow, /.■ 349, 

nom. niid. 
iq26. Mustela punctata Domaniewski, Ann. Mus. Zool. Polon. H.N. 5.' 55. Darasun, 

Eastern Transbaikalia. 
1938. Mustela rixosa pygmaea G. Allen, Manim. Cihina cS: Mongolia, /.• 383. 
Range: Eastern .Siberia, Manchuria, Mongolia. 

Mustela nivalis g.^linthi.\s Bate, 1906 

1906. Putorius nivalis galinthias Bate, P.Z.S. /poj, 2: 319. Crete. (Listed as a distinct 

species alliecl to ''africana" = numidica by Miller (1912).) 

Mustela niv.^lis dinniki Satunin, 1907 

1907. Putorius nivalis dinniki Satunin, Mitt. Kaukas. Mus. Tiflis, jj.- 105 (Russian), 

151 (German). .Sta\Topol, Caucasus. 

Mustela nivalis russelliana Thomas, 191 1 

191 1. Mustela russelliana Thomas, Abstr. P.Z.S. 4; P.Z.S. 168. Tatsienlu, Szechuan, 

China. G. Allen ( 1938) retains this as a species, suggesting it is near stoliczkana. 

Based on one adult female and three other immature specimens. 

MusTEL.\ Niv.^Lis NAMiYEi Kuroda, 1 92 1 

1921. Mustela rixosa namivei Kuroda, J. Mamm. r.- 209. Awomori, Northern Hondo, 

(?) 1936. Mustela pygmaea resoid'una Kishida, Dobuts Zasshi. ^8, 4: 177. Hokkaido, 

(?) 1936. Mustela prgniaea caraftensis Kishida, loc. eit. S.ikhalin. 
Ranges to the Kuriles. 

Mustela nivalis mosanensis Mori, 1927 

1927. Mustela nivalis mosanensis Mori, J. Clhosen N.H. Soc. §: 28. Vengan, near 
Mosan, Korea. 

Mustela nivalis trett.^ui Kleinschmidt, 1937 

1937. Mustela trettaui Kleinschmidt, Falco, 23- ■ ■• Germany. 


MusTELA (?) NIVALIS TONKiNENSis Bjorkegren, 1942 

1942. Mustela tonkinensis Bjorkegren, Ark. Zool. jjjB, 15: i. Chapa, Tonkin, Indo- 

Mustela altaica group 

Mustela altaica Pallas, 181 1 Alpine Weasel 

Approximate distribution of species: in the U.S.S.R., from Ussuri region west- 
wards to region of Lake Baikal, Altai Mountains, and mountains of Eastern Russian 
Turkestan (Tarbagatai, Balkash region, Tianshan, Pamir) ; Mongolia, Manchuria 
and Western Sinkiang (Ognev); Tibet; states of Kansu, Szechuan and Shansi, in 
China; Himalayas, from Kashmir to Sikkim. 

Mustela altaica altaica Pallas, 1 8 11 

181 1. Mustela altaica Pallas, Zoogr. Ross. As. 98. Altai Mountains. 

1823. Putorius alpinus Gebler, Mem. Soc. Imp. Nat. Mosc. 6: 212. Mines of Liddersk, 

Altai Mountains. 
1914. Mustela sacana Thomas, Ann. Mag. N.H. /j.' 566. Near Przewalsk, Djarkent, 

Semirechyia, Eastern Russian Turkestan. 
Range: Siberia and China, as listed under the species. G. Allen recognizes only this 
form in China, but Pocock thought the next race occurred in Tibet, Kansu and 

Mustela altaica temon Hodgson, 1857 

1857. Mustela temon Hodgson, J. Asiat. Soc. Bengal, 26: 207. Sikkim. 
( ?) 1872. Putorius astutus Milne-Edwards, Nouv. Arch. Mus. H.N. Paris, 7, Bull. : 92. 

Moupin, Szechuan. G. Allen thinks this name is a synonym of A/, kathiah. 

Against this opinion see Pocock, 1941, 353 (footnote). 
191 1. Mustela longstaffi Wroughton, J. Bombay N.H. Soc. 20: 931. Teza, Upper 

Sutlej Valley, Northern India, 14,000 ft. 
Range: Himalayas (Sikkim to Gilgit and Karakorum Mountains) and Tibet. 

Mustela altaica raddei Ognev, 1928 

1928. Kolonocus alpinus raddei Ognev, Mem. Sect. Zool. Soc. Amis. Sci. Nat. Moscou, 
No. 2 : 9, 28. Kulusytaevsk village, near Tareinor, Transbaikalia. 

Mustela altaica birulai Ognev, 1928 

1928. Kolonocus alpinus birulai Ognev, Mem. Sect. Zool. Soc. Amis. Sci. Nat. Moscou, 
No. 2: 10, 28. Liangar, Western Pamir Mountains. 

Mustela kathiah Hodgson, 1835 Yellow-bellied Weasel 

Approximate distribution of species: Karakorum Mountains; Kumaon and Nepal, 
eastwards to Assam, Burma, Indo-China; Hupeh, ? Szechuan, Yunnan, Kwantung 
and Fukien in Southern China. 



1835. Miistela (Piitorius) kathiah Hodgson, J. Asiat. Soc. Bengal, 4: 702. The Kachar, 

northern region of Nepal. 
1837. Muitda (Putorius) auriventer Hodgson, J. Asiat. Soc. Bengal, 6: 563. 
1895. Piiloriiis doisalisTroueiiS3.Tt, Bull. Mns. H.N. Paris, /,• 235. Tatsienlu, Szechuan, 

(?) iqio. Arclogalt tsaidamensis Hilzheimer, Zcxil. Anz. jjj.' 309. Tsaidam Mountains, 

\Vestern Kokonor. G. Allen ( 1938, 380) suggests this is the same -as either 

kathiah or altaica, it is not clear which. 
1922. Arclogalc rtuili Matschie, Arch. Nat. 88, Sect. A, 10: 17. Kwantung, Southern 


MusTEL.\ K.\THiAii CAPORiAccoi de Bcaux, 1935 

1935. Muslfla kathiah cajioriaccoi de Beau.x, Atti Soc. Ligust. 14: 65. Baltoro, Kara- 
korum Mountains, Kashmir. 

Muitiia siliirica group 

(Mustela sihirica is type oi' h'olonokus Satunin, if further subgeneric division of the 
genus is required.) 

Mustela sibirica Pallas, 1773 Siberian Weasel 

Appro.ximate distribution of species: In the U.S.S.R., "whole of the forest part of 
Siberia north approximately to the limit of the full-grown forest, and south-west to 
the Altai and adjoining areas, inclusive. Does not occur in Kamtchatka, the Shantar 
Islands and Sakhalin. \Vest of the Ural range it extends as far as Bashkiria, the ad- 
joining part of Cihkalovsk Province, the eastern half of Tatary and Kirov Pro\-ince" 
(Bobrinskii). Japan, Formosa, and throughout China, Manchuria, Tibet. Himalayan 
India, fmni Kashmir eastwards to Northern Burma. Ja\a. 

MlSTF.LA SIBIRICA siEiRi<:.\ Pallas, 1773 

1773. Miistila sihirica Pallas, Reise. Russ. Reichs. ;■, appendix: 701. \'orposten 

Tigerazkdi, near Ust-Kamenogorsk, Western Altai. 
(?) 1904. Miistiia iibuica miles Barrett-Hamilton, Ann. Mag. N.H. /j\- 391. Dauria, 

Transbaikalia, Eastern Siberia. 
11)1 I. h'olonokui sihiricus australis Satunin, .Mitt. Kauk. Mus. 5.' 265, 280. Tyumen 

district. Western .Siberia. 
Range: Russia and Siberia as under the species, except the Far East. 

-Mustela sibirica sibhemachal.\n.'\ Hodgson, 1837 

1037. Mustila 'Piitoruis] uilihtinaiiialaNin Hi)d<,'s(inv ]. .\siat. Soc. Bengal, 6: 563. 

ic'!j2. Mintila hinuriiilis Blyth, J. .\siat. .Soc. Bengal, //.' 99, 280 (footnote). .Sikkim. 
1M43. Mustila liiin/ii'lilii Gray, Ann. Mag. N.H. //.■ 118. Bhutan. 
Range: Ne])al In lihutan, 5,000-16,000 ft. 



1842. Mustela canigula Hodgson, J. Asiat. Soc. Bengal, //.■ 279. Lhasa, Tibet. Perhaps 

ranging to Nepal. 

Mustela sibirica hodgsoni Gray, 1843 

1843. Mustela hodgsoni Gray, Ann. Mag. N.H. //.• 1 18. Himalayas. Range: Kashmir 

and Western Himalayas, from Chamba to Garwhal, 7,000-9,000 ft. 

Mustela sibirica itatsi Temminck, 1844 

1844. Mustela itatsi Temminck, Fauna Japonica, Mamm. 34, pi. vii, fig. 2. Japan. 
1844. Mustela natsi Temminck, op. cit. 34 (footnote). This form is tentatively included 

as a race of j\/. sibirica on the basis of the B.M. material. 
Range: Hokkaido, Hondo, Shikoku, Kiushiu, Iki Island, Japan. 

Mustela sibirica davidiana Milne-Edwards, 1872 

1872. Putorius davidianus Milne-Edwards, Nouv. Arch. Mus. H.N. Paris, 7, Bull.: 92 

(footnote). Kiangsi, Southern China. 
1904. Putorius sibiricus noctis Barrett-Hamilton, Ann. Mag. N.H. /j.- 390. Sanyentze, 

Fukien, South-Eastern China. 
1913. Mustela (Lutreola) taivana Thomas, Ann. Mag. N.H. 12: 91. Mt. Arizan, 

8,000 ft. Formosa. (For status, see Pocock, 1941, 370.) 
1922. Lutreola melli Matschie, Arch. Nat. 88, Sect. A, 10: 35. Canton region. Southern 


Range: South-Eastern China, north to Hupeh, and Formosa. 

Mustela sibirica fontanieri Milne-Edwards, 1871 

1871. Putorius fonlatiierii Milne Edwards, Rech. Mamm. 205, pi. 61, fig. i. Peiping 
(Pckin), China. 

1907. Lutreola stegmarini Matschie, ^Viss. Ergebn. Exped. Filchner to China, 10, i : 

150. Near Tsingtao, Shantung, China. 

Range: Northern China, Shantung, Chihli, Shensi and Shansi. 

Mustela sibirica moupinensis Milne-Edwards, 1874 

1874. Putorius moupinensis Milne-Edwards, Rech. Mamm. 347, pis. 59 (fig. 2) and 60 

(fig. 4). Moupin, Szechuan, China. 
1910. Lutreola major Hilzheimer, Zool. Anz. j§: 310. Near Sungpan, Northern 

Szechuan, China. Not of Fatio, 1905; nor Nilsson, 1820. 
191 o. Lutreola tafeli Hilzheimer, loc. cit. Near Sungpan, Szechuan, China. 
1 92 1. Mustela hamptoni Thomas, J. Bombay N.H. Soc. 2j: 500. Mt. Imaw-bum, 

Kachin Province, 9,000 ft.. Northern Burma. 

Range: Szechuan, Kansu, Yunnan and Northern Burma. 

Mustela sibirica quELP.^RTis Thomas, 1908 

1908. Lutreola quelpartis Thomas, P.Z.S. 53. Quelpart Island, Korea. 


palae,\rc:tic: and indian mammals 1758-1946 


igii. MiisUia manchurica Brass, Reiche Pelze, 490. Manchuria. 

1 93 1. Kolonocus sibiricus katsurai Kishida, Dobuts Zasshi. 4;): 380, nom. nud. 

Range includes the Far East of Siberia. 

MusTELA SIBIRICA SHO Kuroda, 1924 

1924. Lutnola ilatsi sho Kuroda, on New Mamm. from Riu Kiu Islands and vicinity, 

Tokyo, 10. Miyanoura, Yakushima Island, Japan. Range: Tanegashima 

and Yakushima, south of Japan. 

MusTELA SIBIRICA coREANA Domauiewski, 1926 

1926. Kolonocus sibiricus coreanus Domaniewski, Ann. Zool. Mus. Polon. 5.- 55. Seoul, 

1 93 1. Kolonocus sibiricus peninsulae Kishida, Dobuts Zasshi. 4;^: 380, nom. nud. 

MusTELA SIBIRICA CHARBiNENSis Lowkashkin, 1934 

1934. -^ii'^l'l" [Kolonocus] sibirica charbinensis Lowkashkin, China J. Sci. & Arts, 20: 
40. Krcstowsky Island, in Sungai River, near Harbin, Manchuria. 


1943. Muslela sibirica asaii Kuroda, Bull. Biogeogr. Soc. Tokyo, ij, 8: 55. Oshima 
Island, Izu Islands, Japan. 

Muslela lutreola group 
For this group, Lutreola Wagner, 1841, is available. It is given subgeneric rank 
by many authors. 

Mustela lutreola Linnaeus, 17G1 European Mink 

Approximate distribution of species: from Western France, eastward to the Tobol 
and Irtish Rivers in Western Siberia; south to Austria, Hungary, Rumania and 
Transcaucasia; north to Finland and Northern Russia (Harper, 1945). (Bobrinskii 
states it ranges to Northern Caucasus only, not Transcaucasia, and quotes it also 
from Yugoslavia and Italy.) Distribution includes Poland. 

Mustela lutreola lutreola Linnaeus, 1761 

I 761. Viverra lutreola Linnaeus, Faun. Succ. 5. Finland. 

1777. Lutra minor Er;<lcben, Syst. Regn. Anini. /.• 4-,:. Renaming cA lutreola. 

1792. Mustela Lutra fulva Kerr, Anim. Kingd. 173. Renaming ui lutreola. 

(?) 1839. Mustela lutreola var. alba de Selys Longchamps, Etudes Micromamm. 46, 

nom. nud. 
(?) 1B63. Putorius alpinus Ogcrien, H.N. du Jura, j.- 59. Highest portions ofjura. Not 

ofGeblcr, 1823. 
1879. Lutreola europaea Homeyer, Zool. Garten, 20: 184. Substitute for lutreola. 
1912. MuUela (Lutreola) lutreola ivvborgensis Matschie, S.B. Gcs. Nat. Fr. Berlin, 347. 

\'iborg, Finland. 
Range: according to Bobrinskii, Finland, northern part of Russia as far south as 
Leningrad Province, Gorki, Sverdlovsk, possibly Bashkiria. 



191 2. Muslela {Lutreola) lutreola cylipena Matschie, S.B. Ges. Nat. Fr. Berlin, 348. 

East Prussia. 
191 2. Mustela [Lutreola] lutreola budina Matschie, loc. cit. 349. Ortelsburg, East Prussia. 
1912. Mustela (Lutreola) lutreola varina Matschie, loc. cit. 351. Schwerin, Mecklenburg, 

1 91 2. Mustela {Lutreola) lutreola albica Matschie, loc. cit. 351. River Levitz, tributary 

of Elbe, Mecklenburg, Germany. 
1 91 2. Mustela [Lutreola] lutreola glogeri Matschie, loc. cit. 354. Brieg, Silesia. 
Range: Lat\ia, Lithuania, Germany, ? Western White Russia. 

Mustela lutreola biedermanni Matschie, 191 2 

1912. Mustela [Lutreola] lutreola biedermanni Matschie, S.B. Ges. Nat. Fr. Berlin, 353. 

Malicorne, South-\Vestern France. 
1912. Mustela [Lutreola) lutreola aremorica Matschie, loc. cit. 354. Near Vimont, Caen, 


Mustela lutreola transsylvanica Ehik, 1932 

1932. Mustela lutreola transsylvanica Ehik, Allat. Kozlem, 2g: 142. Kovaszna, Tran- 

1932. Mustela lutreola hungarica Ehik, Allat Kozlem, 2g: 142. Komitate Turoc 
Hungary. Not Mustela eversmanni hungarica Ehik, 1928. 

Bobrinskii only recognizes one race from Hungary, which he quotes under the 
preoccupied name hungarica from Hungary, Rumania, Southern Germany, Yugo- 
slavia, Italy, apparently Bessarabia. 

Mustela lutreola turovi "Kuznetzov & Novikov," 1939, Bobrinskii, 1944 
1944. L[utreola) l{utreola) turovi Bobrinskii, Mamm. U.S.S.R. 127. No exact locality, 

'"The Caucasus mink; distributed south of the proceeding form" (= M. I. 

borealis, here renamed novikovi). ^Ve are unable to trace any other reference 

to this form than that given here. 

Mustela lutreola novikovi nom. nov. 

1939. Lutreola lutreola borealis Noviko\-, The European Mink (Leningrad), 63. Valley 

of the River Byonki, near Milet, Bogorodsk region, Moscow Govt., Russia. 

Range: Estonia, Eastern Latvia, White Russia, across central zone of 

European U.S.S.R. to Bashkiria, apparently as far south as the forest-steppe 

zone. Not Mustela jiavigula var. borealis Radde, 1862. 

Mustela lutreola binominata nom. nov. 

1939. Lutreola lutreola caucasica Novikov, The European Mink (Leningrad), 63. 
Station Prochladnaya, Northern Caucasus. Not of Barrett-Hamilton, 1900. 

Alustela strigidorsa group 

Referrable to Pocockictis Kretzoi, 1947, if further subgeneric division of the genus 
is required. 


i'ai.aearctk: axd Indian mammals i7-,8-i94r, 

Mustela strigidorsa Gia\-, 1853 Back-striped Weasel 

Approximate distribiitidii of species: Nepal, Sikkini, Assam, Burma, Tcnasscrini 
and Indo-China. 

Mustela strigidorsa Gray, 1853 

i8-,3. Miislela stri^odorsa Gray (Hodgson MS.), P.Z.S. igi. Sikkim. 
i8r,5. Mustela ^lrii;.idorsa Hnrsficid, Ann. Mag. N.H. 16: 107. 
Range : as abov c. 

Suhgciuis PUTORIUS Cinier, 181 7 

Mustela putorius Linnaeus, 1758. European I^)lrcat 

Approximate distribution of species (as understood by Pocock. 1936): Britain, 
Norway, Sweden, Holkand, Germany, France, Belgium, Denmark, Poland, Switzer- 
land, south to Spain, Italy, Rumania; Finland. The greater part of Russia, north to 
^Vhite Sea, south to Crimea, Northern Caucasus, etc., Kazakstan and Southern 
Siberia as far cast as the Amur region, approximately. Mongolia, and Palaearctic 
parts of China (southwards about to Szechuan); Tibet; Kashmir; Palestine, accord- 
ing to Bodenheimcr; Morocco. The distribution of the Ferret is of course subject to 
modification by human agency. 


1758. Miistda putorius Linnaeus, Syst. Nat. loth cd. /: 46. Sweden. 

1 78-). Mustela litis Boddaert, Elench. Anim. 87. Renaming oi^ putorius. 

1795. Mustela furo-pulorius Link, Beytr. Naturgesch. /.■ 83. 

1798. Viverra foetens Thunberg, Bcskrifning pa Svenske Djur, i -,. Renaming of 

1801. Mustela putorius albus Bechstein, Gcmeinn. Dcutschlands, 2nd ed. /.•782. 

Thuringia, Germany. Not alba, he. cil. "]-)<>. 
1827. Putorius vulgmrs Grifhth, Cuvier's Anim. Kingd. j: 120. Renaming of 

(?) 1839. Mustela putorius flavieans dc Sclys Longchamps, Etudes de Micromamm. 

145, uom. nud. 
(?) 1839. Mustela putorius \ar. vison de Sclys Longchamps, loc. eit., nam. nud. 
1843. Putorius Joetidus Gray, List. Spec. Mamm. B.M. 64. Renaming oi^ putorius. 
1851. Putorius verus Simashko, Russ. Fauna, 2: 357. 

1863. Putorius iiifectus Ogcricn, H.N. du Jura, 3: 59. Substitute i'nv putorius. 
1904. Putorius putorius manium Barrett-Hamilton, Ann. XLag. N.H. / j.- 390. Teufin, 

Apfenzell, Switzerland. 
i()2(3. Putorius putorius stantsehinskii Mehmclcr, Wiss. .Mitt. Univ. Smolensk, 137. 

Smr)lensk Govt., Russia. 
(?) 1929. Mustela putorius orientalis Brauncr, Ukr. Misl. ta Ribalka, 2-3, 8-9. No 

locality. Not of Ognev, 1928. (.Y.r.) 

Range: Europe, from Scandina\ia to Northern Spain and Mediterranean coast, 
westwards to Britain, eastwards to the Ural Mountains. 



MusTELA puTORius FURO Linnaeus, 1758. Ferret 

1758. Mustela furo Linnaeus, Syst. Nat. loth ed. /; 46. "Africa." 
1865. Pittorius foetidus var. subrufo Gray, P.Z.S. no. Bred in captivity. See page 252 
for discussion and status. 

Mustela putorius eversmanni Lesson, 1827 

1827. Mustela eversmanni Lesson, Man. de Mamm. 144. Between Orenburg and 
Bokhara, Russian Turkestan. 

1842. Mustela putorinus 'Qlyth, ]. Asiat. Soc. Bengal, //, i: 281. 

(?) 1944. M(uslela) ev{ersmanm) satunini "Migulin, 1928," Bobrinskii, Mamm. 
U.S.S.R. 126. Nagaiskie steppes. We are unable to trace the reference from 
Migulin, 1928. Bobrinskii treats M. eversmanni as a distinct species. 

Mustela putorius larvatus Hodgson, 1849 

1849. Putorius larvatus Hodgson, J. Asiat. .Soc. Bengal, 18: 447. Utsang, north of 

Sikkim, in Southern Tibet. 
1851. Putorius tibetanus Horsfield, Cat. Mamm. E. Ind. Co. 105. Utsang, Southern 

Range: Tibet and Kashmir. Bobrinskii considers this as a subspecies of eversmanni. 

Mustela putorius aureola Barrett-Hamilton, 1904 

1904. Putorius putorius aureolus Barrett-Hamilton, Ann. ^L^g. X.H. ij: 389. Ferrol, 
Province of Coruiia, Spain. 

Mustela putorius michnoi Kastschenko, 19 10 

1910. Putorius eversmanni var. michnoi Kastschenko, Ann. Mus. Zool. Acad. Sci. St. 

Petcrsb. ij: 271. River Kiran, 20 km. from Troizkosavsk, Transbaikalia. 
1913. Mustela lineiventer Hollister, Proc. Biol. Soc. Washington, 26: 2. Tchegan- 

Burgazi Pass, Little Altai, Siberia. 
Range: Transbaikal steppes, according to Bobrinskii, who thinks it may be the same 
as larvatus and regards it as a subspecies oi eversmanni. 

Mustela putorius tiarata Hollister, 1913 

1913. Mustela tiarata Hollister, Proc. Biol. Soc. Washington, 26: 2. Chiuningchow, 
150 miles east of Lanchow, Kansu, China. Range: Mongolia, Kansu, 
Shansi, Szechuan. Treated as a subspecies oi eversmanni by G. Allen. 

Mustela putorius talassica Ognev, 1928 

1928. Putorius eversmanni talassicus Ognev, Mem. Sect. Zool. Soc. Amis. Sci. Nat. 
Moscou, 2: 26, 30. Talassky Alatau (north-east of Tashkent), Russian 
Turkestan. (Bobrinskii gives Dzhinak Golodnaya Steppe as the locality.) 

Mustela putorius hungarica Ehik, 1928 

1928. Mustela eversmanni hungarica Ehik, Ann. H.N. Mus. Hung. 2§: 37. Magyarova, 

(?) 1944. .M(ustela) ev(ersmanni) occidentalis "Brauner, 1929," Bobrinskii, \Limm. 

U.S.S.R. 126. Former Kherson Govt., Russia. W^e are unable to trace 

reference from Brauner, 1929. 


pai-aearc:tio and Indian mammals 1 7^,8-1946 

MrsTELA puTORU's AMURENSis Ognev, 1930 

IQ30. Piitorius evenmanni amurerisis Ognev, Okhntnik, No. ii: 25. Blagosveschcnsk 
region of Amur Basin, Eastern Siberia. 


1932. MuUfla putorius rotkschildi Pocock, .Scot. Nat. Edinb. 103. Malcoci, Dobrudscha, 


1936. Pulorius pu/orius angiitis Pocock, P.Z.S. G94. Liangammarch, Brecknockshire, 


1936. Piitorius pulorius aureus Pocock, P.Z.S. 703. Kazan, Central Russia. 


1936. Pulorius pulorius adrniralui Pocock, P.Z.S. 706. Chihfeng, Chihii, North-Eastern 


1939. Pulorius pulorius caledoniae Tetley, P.Z.S. Ser. B., 37. Lochinver, Sutherland, 

Incerlae sedis 

Mustda rasarhdyi Krctzoi, 1942, Eoldt. Kozl. Budapest, jr.- 349, new name for: 
Muslda hungarica Vasarhelyi, 1942, Zool. Anz. Leipzig, /jj/.- 221-226; not of Ehik, 

1929 {M. eversmanni hungarica) nor of Ehik, 1932 (M. lulrcola hungarica). 

Hungary. (N.V.) 

Genus VORMELA Blasius, 1884 

1884. ]'ormeIa Blasius, Bericht der Naturforsch. Gescllsch. in Bembcrg, /j.- 9. Mustela 
sarmatica Pallas = Muileia pcrrgusna Guldenstaedt. 

I species: I'onnela pcrcgiisna, page 266 

Vormela peregusna Guldenstaedt, 1770 Marbled Polecat 

Approximate distribution of species: Rumania, Bulgaria, Black Sea steppes, 
Crimea, Ciscaucasia, and Kazakstan to Western Altai foothills, Transcaucasia; Asia 
Minor, Palestine, Syria, Iraq, Persia, Afghanistan; Baluchistan; Mongolia. 

VOR.MEI.A PEREGUSNA PERF.GUS.N'.-\ Gtildcnstacdt, 1770 

1770. Minlcla pert-gusna Guldenstaedt, Nov. Comm. Acad. Sci. Imp. Pctrop. 14, i: 

441. Banks of the River Don, Southern Russia. 
I 77 I. Muslda sainialica Pallas, Reise Prov. Russ. Reichs, /.• 453. Along the Volga 

River, Southern Russia. ^\ccording to Chaworth-Musters, Sysran; text, 

Inc. cil. I.- 175.) 



1935. Vormela peregusna peregusna natio intermedia Ognev, Mamm. E. Europe, N. Asia, 

5; 70. Village Starogradskaia, River Terek, Kisljar subdistrict, Terek 
district, Caucasus. 
Range: eastwards to Western Siberia. 

Vormela peregusna alpherakii Birula, 1910 

1910. Vormela sarmatica alpherakii Birula, Ann. Mus. Zool. St. Petersb. i§: 333. Trans- 

caspia, near Ashabad. 
1910. Vormela koshewnikowi Satunin, Zool. Anz. 36: 59. Ashabad, near Persian 

border, Russian Turkestan. 
1910. Vormela tedshenika Satunin, Zool. Anz. 36: 60. Tejen Oasis, "Oase Tedschen," 

Russian Turkestan. 
Range : Russian Turkestan, part, Persia, Afghanistan, Baluchistan. 

Vormela peregusna negans Miller, 1910 

1910. Vormela negans Miller, Proc. U.S. Nat. Mus. 38: 385, pi. 17. Ordos Desert 

(about 100 miles north of Yulinfu, Northern Shensi), Inner Mongolia. 

Ranges into Eastern Turkmenia, according to Bobrinskii. 

Vormela peregusna euxina Pocock, 1936 

1936. Vormela peregusna euxina Pocock, P.Z.S. 718. Malcoci, Dobrudsha, Rumania. 

Vormela peregusna syriaca Pocock, 1936 

1936. Vormela peregusna syriaca Pocock, P.Z.S. 720. Tiberias, Syria. Range: to Western 
Iraq. (Specimens in B.M. (of this race?) from Palestine.) 

Vormela peregusna ornata Pocock, 1936 

1936. Vormela peregusna ornata Pocock, P.Z.S. 721. Neighbourhood of Lake Baikal, 

Genus POECILICTIS Thomas & Hinton, 1920 

1920. Poecilictis Thomas & Hinton, Ann. Mag. N.H. 5.- 367. Mustela libyca Hemp- 
rich & Ehrenberg. 

I species: Poecilictis libyca, page 267 

Poecilictis libyca Hemprich & Ehrenberg, 1833 Libyan Striped Weasel 

Approximate distribution of species : Northern Africa, from Morocco and Algeria 
to Libya and Egypt, south to the Sudan and Northern Nigeria. 

Some earlier authors, Trouessart included, quoted this species from Asiatic Turkey, 
but we have not been able to verify it as occurring in any part of Asia, and suspect 
these allusions were caused by confusion with some other small striped Mustelid. 

Poecilictis libyca libyca Hemprich & Ehrenberg, 1833 

1833. Mustela libyca Hemprich & Ehrenberg, Symb. Phys. Mamm. 2: k verso. Libya. 
Range includes Lower Egypt. 

s 267 


1856. ^orillii vniUanlii Lochc, Rev. Mag. Zool. 8: 497, pi. 22. Algeria. Range includes 
Tunis and Morocco. 

Subfamily M e 1 1 i v o r i n a e 

Genus MELLFVORA .Storr, 1780 

1780. Mdlivora StuFr, Prodr. Meth. Mamni. 34, and Tab. A, Mamm. Viverra ratel 

Sparrmann = Viverra capensis Schreber. 
1827. Ratellus Gray, Griffith's Cuvier Anini. Kingd. f^: 1 18. Viverra capensis Schreber. 
1836. Ursitaxus Hodgson, .Vsiat. Res. ifj: 61. I'nilaxus inauritus Hodgson. 
1841. .\/(7i7o/)'.v Gloger, Gemein. Nat. /.• 57. Viverra capensis Schreber. (Type selected 

by Pocock, 1941, 454-) 
1843. Lipotus Sundevall, Svcnska \'ct. Ak. Handl. 1842: 199. I'rsus mellivnrus Ckuicr 

= Viverra capensis Schreber. 

I species: Mdlivora capensis, page 268 

Mellivora capensis Schreber, 1776 Ratel, or Honey Badger 

Approximate distribution of species: Southern Turkmenia (River Atrek, Kopet- 
Dag, River Tedshen) ; Syria, Palestine, Afghanistan (according to Bobrinskii), Persia, 
Arabia; India, from North-West Frontier and Nepal, south to Sind, Cutch, Bengal, 
thence to the Madras Presidency. Morocco; Ethiopian Africa from Asben on the 
west, the Sudan, Abyssinia and Somaliland on the cast, southwards to the Transvaal, 
and the Cape Province (Little Namaqualand .uid Uitenhage district). 

(Mellivora capensis capensis Schreber, 1776. E.xtralimital) 

1776. I'iverra capensis Schrcbci', Saugcth. pi. 125, also 1777, j: 451. C'ape of Good 

Mellivor.-\ cape.nsis iNDiCA Kerr, 1792 

1792. Ursus indicus Kerr, Anim. Kingd. 188. India. 

1830. Ratelus mellivorus Bennett, Garclens & Menag. Zool. Soc. Quad i : i ■;. Interior 

of Madras. 
1835. Ratelus indicus Burton, P.Z.S. 113. Upper Provinces of Bengal. 
1851. Mellivora ratel Horsfield, Cat. Mamm. E. Ind. Co. 120. India. 
1862. Mellivora ratelus PVaser, Cat. Z. Gdns. 9. 

Range: Sind, Cutch, Hazaribagh, Western India, to South- Western Russian 

Mellivora capensis inaurita Hodgson, 1836 

1836. I'rsttaxus inauritus Hodgson, .\siat. Res. i<): 61. .Muckuanpur, in Idntliills of 

Southern Nepal. Range: foothills of Southern Nepal, possibly Kumaon and 

North-\\'est Frontier. 



Mellivora capensis leuconota Sclater, 1867 

1867. Mellivora leuconota Sclater, P.Z.S. 98, pi. 8. West Africa. Range: northwards to 
Southern Morocco. 

Mellivora capensis wilsoni Cheesman, 1920 

1920. Mellivora wilsoni Cheesman, J. Bombay N.H. Soc. 2y: 335. Ram Hormuz, 
500 ft., Arabistan, South-Western Persia. 

Mellivora capensis pumilio Pocock, 1946 

1946. Mellivora capensis pumilio Pocock, P.Z.S. ii;j: 314. Hadramaut, Southern 

Subfamily M e 1 i n a e 

Genus MELOGALE I. Geoffroy, 1831 

1831. Melogale I. Geoffroy, Belanger, Voy. Zool. Indes Orient. 129 (19 March). 

Melogale persona/a Geoffroy. 
1831. Helictis Gray, P.Z.S. 94 (5 August). Helictis moschata Gray. Valid as a subgenus. 
1922. Nesictis Thomas, Ann. Mag. N.H. g: 194. Helictis everetti Thomas, from Borneo. 

2 species in the area covered by this list: 
Melogale moschata, page 270 
Melogale personata, page 269 

Some authors, including Pocock, have referred the Ferret-Badgers to the genus 
Helictis Gray, 1831, and either discarded Melogale I. Geoffroy under the impression 
that it dated from 1834, or used it as a subgenus of Helictis. But Geoffroy's name dates 
from 19 March 1831, a few months earlier than Gray's name of 5 August 1831. (For 
the date of publication oi Melogale I. Geoffroy, see Sherborn, 1901, Ann. Mag. N.H. 
j: 390.) Pocock (194 1, 396) gave the characters of the two species and recognized no 
subgenus. Simpson (1945, 1 14) lists both Melogale and Helictis as full genera. We take 
a middle view, and here regard Helictis as a subgenus of Melogale. 

Subgenus MELOGALE I. Geoffroy, 1831 

Melogale personata Geoffroy, 1831 Burmese Ferret-Badger 

Approximate distribution of species: Nepal, Assam, Burma, Siam, Indo-China. 

Melogale personata personata Geoffroy, 1831 

1831. Melogale personata I. Geoffroy, Belanger, Voy. Zool. Indes Orient. 137, pi. 5. 
Near Rangoon, Burma. Ranges to Assam, Manipur. 



Melogale personata nipalensis Hodgson, 1836 

1836. Gulo nipalensis Hodgson, J. Asiat. Soc. Bengal, 5; 237. Nepal. 

1888. Helictis orientalis Blanford, Mamm. Brit. Ind. 173. Not of Horsfield, 1821. 

Range: Nepal to Bhutan Duars. 

Melogale personata pierrei Bonhote, 1903 

1903. Hdictis pierrei Bonhote, Ann. Mag. N.H. 12: 592. Near Saigon, Cochin-China. 

Melog.^le personat.'>i laotum Thomas, 1922 

1922. Melogale personata laotum Thomas, Ann. Mag. N.H. g: 194. Nan, 200 m., .Siam. 
Ranges into Indo-China (part). 

Melogale person,\ta tonquinia Thomas, 1922 

1922. Melogale tonquinia Thomas, Ann. Mag. N.H. g: 195. Yen-bay, Songkoi River, 
Tonkin, North Indo-China. (Osgood (1932) thought this was a synonym of 


Subgenus HELICTIS Gray, 1831 

Melogale moschata Gray, 1831 Chinese Ferret-Badger 

Approximate distribution of species: China, from Szechuan southwards to Yunnan, 
thence to Fukicn and Hainan; Formosa; Assam and Burma; Indo-China. 

Melogale moschata moschata Gray, 1831 

1831. Helictis moschata Gray, P.Z.S. 94. Canton, Kwantung, Southern China. The 
range includes Yunnan and Hainan. 

Melogale moschata subaurantiaca Swinhoe, 1862 

1862. Helictis subaurantiaca Swinhoe, P.Z.S. 355. Formosa. 

1922. Helictis subaurantiaca modesta Thomas, Ann. Mag. N.H. g: 196. Bankoro, 
Formosa. For status, see Pocock (1941, 404). 

Melog.\le moschata ferreogrise.'^ Hilzheimer, 1905 

1905. Helictis ferreo-griseus Hilzheimer, Zool. Anz. 2g: 298. Near Hankow, Hupeh, 
China. Range: Szechuan, Fukien and adjacent states, China. G. Allen 
(1938, 396) lists a specimen from Shansi — "probably not native there". 

Melogale mosch,\ta millsi Thomas, 1922 

1922. Helictis millsi Thomas, J. Bombay N.H. Soc. 28: 432. Mokokchung, Naga 
Hills, 5,000 ft., Assam. Range includes Northern Burma. 

Melog.\le mosch.^ta taxilla Thomas, 1925 

1925. Helictis taxilla Thomas, P.Z.S. 500. Ngai-tio, Tonkin, 3,100 ft., Northern Indo- 



Melogale (?) MoscHATA SORELLA G. Allen, 1929 

1929. Helectis taxilla sorella G. Allen, Amer. Mus. Nov. No. 358, 8. Futsing, Fukien, 
South-Eastern China. Not, we think, "Helictis taxilla sorella", as Pocock 
(194 1, 401) shows that taxilla is very close to, if not identical with, millsi. On 
the other hand, G. Allen (pp. 396, 398) lists specimens of both sorella and 
ferreogrisea from Futsing, Fukien. Possibly, therefore, sorella will have to be 
given specific rank. See also Pocock (1941, 405). 

Genus MELES Brisson, 1762 

1762. Aleles Brisson, Regn. Anim. 13. Ursus meles Linnaeus. Hopwood (1947, P.Z.S. 
533-536) would disregard Brisson and date Meles, with type Ursus meles 
Linnaeus, from Boddaert, 1785, Elench. Anim. /.• 45. 

1795. Taxus Cuvier & Geoffroy, Mag. Encyclop. 2: 187. Ursiis meles Linnaeus. 

1815. Melesium Rafinesque, Anal, de la Nature, 59. Renaming of Taxus. 

1925. Meledes Kastschenko, Bull. Acad. Sci. Ukr. Phys.-Math. /, 4: 21. Meles taxus 
arenarius Satunin. 

I species : Meles meles, page 2 7 1 

Most authors seem now to agree that there is only one valid species in this genus. 

Meles meles Linnaeus, 1758 Badger 

Approximate distribution of species: except that it does not occur in North Africa, 
essentially throughout the Palaearctic region, and in Southern China somewhat 
south of that region. 

(In detail, British Isles, west to Ireland, Norway, Sweden, Belgium, France, 
Holland, Denmark, Germany, ? Switzerland, Hungary (B.M.), Poland, Spain, Italy, 
Crete. Widely distributed in the U.S.S.R., according to Bobrinskii the whole of 
Russia except the Pechora basin; Turkestan and across Siberia approximately as far 
north as a line from Surgut-on-Ob to Nikolaevsk-on-Amur; does not occur in Sak- 
halin; Chinese Turkestan, Tibet, Mongolia, Korea, Japan. Throughout the main 
states of China, except, evidently, Yunnan. Asia Minor, Persia and Palestine. (Tate, 
1947, quotes M. m. leucurus from extreme Northern Burma.) ) 

Meles meles meles Linnaeus, 1758 

1758. Ursus meles Linnaeus, Syst. Nat. loth ed. /.• 48. Upsala, Sweden. 

1785. Meles taxus Boddaert, Elench. Anim. /.■ 80. Europe. 

1788. Ursus meles alba Gmelin, Syst. Nat. 13th ed. /.• 102. 

1788. Ursus meles maculata Gmelin, loc. cit. 

1808. Taxus vulgaris Tiedemann, Zoologie, /." 376. Renaming of Ursus meles. 

1816. Meles europaeus Desmarest, Nouv. Diet. H.N. 3: 465. Renaming of meles. 
1827. Meles communis Billberg, Synop. Faun. Scandinaviae, 16. Renaming of meles. 
1827. Afeles communis caninus Billberg, loc. cit. 17. Scandinavia. 

1899. Aleles meles typicus Barrett-Hamilton, Ann. Mag. N.H. 4: 384. 

1906. Aleles meles britannicus Satunin, Mitt. Kauk. Mus. 2: 115. Based on cranial 

measurements of English specimens recorded by Barrett-Hamilton, 1899, 

Ann. Mag. N.H. 4: 384. 
Range: from Italy, north to Scandinavia, west to Ireland, east to Russia. 



Meles meles anakuma Temminck, 1844 

1844. Meles anakuma Temminck, Fauna Japonica, Mamm. 30, pi. 6. Environs of 

Nagasaki and Awa, Japan. Range: Hondo, Shikoku, Kiusiu, ? Hokkaido, 


Meles meles leucurus Hodgson, 1847 

1847. Taxidea leueurus Hodgson, J. Asiat. Soc. Bengal, iG: 763, pi. 29. Lhasa, Tibet. 

Meles meles amurensis Schrenck, 1859 

1859. Meles taxus amurensis Schrenck, Reisen Amur-Lande, 17, pi. i, fig. i. Amur 

region, not far from mouth of Ussuri River. 
1891. Meles sehrenkii Nehring, S.B. Ges. Nat. Fr. Berlin, 103. Alternative name for 

Range: Amur-Ussuri region, Manchuria. 

Meles meles leptorhynchus Milne-Edwards, 18G7 

1867. Meles leptorhynchus Milne-Edwards, Ann. Sci. Nat. Zool. 8: 374. Near Pekin, 

Chihli, China. 

1868. Meles chinensis Gray, P.Z.S. 207. Amoy, Clhina. 

1907. Ateles hanensis Matschic, Wiss. Ergebn. Exped. Filchner to China, 10, i : 138. 

Hinganfu, Shensi, China. 
1907. Meles sinins^ensis Matschie, he. (it. Siningfu, Kansu, China. 
1907. Meles tsingtauensis Matschie, loc. cit. 142. Tsingtao, Shantung, China. 
Range: China, including states of Chihli, Shantung, Chekiang, Kiangsu, Shansi, 
Shensi, Hunan, Fukien, Kansu, Szcchuan. 

Meles meles canescens Blanford, 1875 

1875. Mdes canescens Blanford, Ann. Mag. N.H. 16: 310. Abadeh, between Shiraz 
and Isfahan, 7,000 ft., Persia. 

Meles meles arenarius Satunin, 1895 

1895. Meles taxus arenarius Satunin, Arch. Nat. /; i i i. Ryn Peski, Astrakhan Govt., 
South-Eastern Russia. Range: Caucasus steppes. 

Meles meles marianensis Graells, 1897 

1897. Meles taxus var. marianensis Graells, Mem. Real. Acad. Cien. Madrid, ij: 170. 
Central Spain. 

1899. Meles meles mediterraneus Barrett-Hamilton, Ann. Mag. N.H. 4: 384. Seville, 


MeLes meles siBiRicus Kastschenko, 1900 

1900. Meles taxus sibincus Kastschenko, Key to Mamm. Tomsk, table 15 (Russia), 

and 1 90 1, Ann. Mus. Zool. Acad. Sci. St. Petersb. 6: 611. Plains of central 
part of Tomsk Govt., Siberia. 

Meles meles raddei Kastschenko, 1901 

1 90 1. Meles amurensis raddei Kastschenko, Ann. Mus. Zool. Acad. Sci. St. Petersb. 6: 

613. Steppes of Transbaikalia, Eastern Siberia. 


Meles meles altaicus Kastschenko, igoi 

igoi. Meles amurensis altaicus Kastschenko, Ann. Mus. Zool. Acad. Sci. St. Petersb. 
6: 613. Coast of Lake Telezkoi, South-Western Russian Altai. 

Meles meles minor Satunin, 1905 

1905. Meles meles minor Satunin, Priroda i. Ochota, 2: 467. (M.V.) 1905, Mitt. 
Kaukas Mus. 2: 113 (German, 288). Borzom, Gouv. Tiflis, Transcaucasia. 

Meles meles arcalus Miller, 1907 

1907. Meles arcalus Miller, Ann. Mag. N.H. 20: 394. Lassethe Plain, Crete. 

Meles meles blanfordi Matschie, 1907 

1907. Meles blanfordi Matschie, Wiss. Ergebn. Filchner E.xped. to China, 10, i : 143. 
Kashgar, Chinese Turkestan. 

Meles meles tianschanensis Hoyningen-Huene, 19 10 

1910. Aleles tianschanensis Hoyningen-Huene, Zur. Biol. Estlandisch. Dachses, 63. 
Tianshan Mountains. 

Meles meles melanogenys J. Allen, 191 3 

1913. Meles melanogenys }. Allen, Bull. Amer. Mus. N.H. ^2.' 433. Musan, Northern 


Meles meles rhodius Festa, 1914 

1914. Meles meles rhodius Festa, Boll- Mus. Zool. Anat. Comp. Torino, 2g: 6. Koskino, 

Island of Rhodes, Eastern Mediterranean. 

Meles meles ponticus Blackler, 1916 

1916. Meles meles ponticus Blackler, Ann. Mag. N.H. 18: 75. Scalita, near Trebizond, 
3,000 ft., Asia Minor. 

Meles meles caucasicus Ognev, 1926 

1926. Meles meles caucasicus Ognev, Bull. Sci. Inst. Expl. Caucasus, /.• 50, 56. Near 
Vladikavkaz (Ordzhonikidze), Caucasus. 

Meles meles tauricus Ognev, 1926 

1926. Meles meles tauricus Ognev, Bull. Sci. Inst. Expl. Caucasus, /.• 51, 56. Chatyr- 
Dag, Beshuisk Forest, Crimea, Southern Russia. 

Meles meles talassicus Ognev, 1931 

1 93 1. Meles leptorhjnchus talassicus Ognev, Mamm. E. Europe, 2: 478. Southern slopes 
of Talasski Alatau, north-east of Tashkent, Russian Turkestan. 

Meles meles hept.neri Ognev, 1931 

1 93 1. Meles meles heptneri Ognev, Mamm. E. Europe, 2: 775. Village of Aleksandro 
Nevskaia, 18 km. north-west of Kisljar, Daghestan, Caucasus. 



Meles meles DANicis Dcgcibol, 1933 

1933. Meles meles danicus Degerbol, Danmarks Pattcdyr i Fortidcn, 574, 634. 
SjacUand, Denmark. 

Meles meles severzovi Hcptner, 1940 

1940. Meles meles severzovi Heptner, Z. Sauget. /j; 224. Region of Arkit, Chodscha- 

ata Valley, south of Tschatkal Mountains, near Lake Sarytschilek, Russian 


Genus ARCTONYX F. Cuvicr, 1825 

1825. Arctomx F. Cuvier, H.N. Mamm. j, pt. 51, pi. and text. Arctonyx collaris Cuvier. 
1891. Trichomanis Hubrecht, Notes Leyd. Mus. 13: 241 . Trichomanis hoevenii Hubrecht 
(the Sumatran race of .-1. collaris). 

I species: Arctonyx collaris, page 274 

Arctonyx collaris F. Cuvier, 1825 Hog-Badger 

Approximate distribution of species: all the larger states of China; Sikkim Terai to 
Assam and Burma; Indo-China, Siam (south at least to Trang) and Sumatra. 

Arctonyx collaris collaris F. Cuvier, 1825 

1825. Arctonyx collaris F. Cuvier, H.N. Mamm. j, pt. 51, pi. and text. Bhutan Duars, 

Eastern Himalayas. 
1853. Arctonyx laxoides Blyth, J. Asiat. Soc. Bengal, 22: 591. Assam. 
1856. Arctonyx isonyx Horsfield (Hodgson MS.), P.Z.S. 398. Sikkim Terai. 
1863. Arctonyx collaris taraiyensis Gray, Cat. Hodgson's Coll. B.M., 2nd ed. 7. Sikkim 

Range: Sikkim Terai, Bhutan Duars, Assam. 

Arctonyx collaris albogularis Blyth, 1853 

1853. Meles albogularii Blyth, J. Asiat. Soc. Bengal, 22: 590. Eastern Tibet. (More 

likely, perhaps, from Szechuan, China?) 
1 87 1. Meles (Arctonyx) obscurus Milne-Edwards, Rech. H.N. Mamm. 200, 202. 

Szechuan, China. 
191 I. Arctonyx leucolaemus orestes Thomas, Abstr. P.Z.S. 27; P.Z.S. 688. Tsingling 

Mountains, Shensi, 12,000 ft., China. 

1922. Arctonyx obscurus incultus Thomas, Ann. Mag. N.H. 10: 395. Chinteh, Anhwei 

(about 150 km. west of Hangchow), China. 
Range: Southern China, northwards to Shensi. For status of this race (which G. 
.Allen thought was a synonym of the typical race) sec Pocock (1941, 427, 434). 

.\rctony.\ collaris leucolaemus Milne-Edwards, 1867 

1867. Meles leucolaemus Milne-Edwards, Ann. Sci. Nat. Zool. 8: 374. Near Pckin, 
Chihli, China. 

1923. Arctonyx leucolaemus milne-edwardsii Lonnberg, Ann. Mag. N.H. //.• 322. Min- 

shan. Southern Kansu, China. 



Arctonyx collaris dictator Thomas, 19 lo 

1910 Arctonyx dictator Thomas, Ann. Mag. N.H. 5." 424. Lamra, Trang, Lower Siam. 
(?) 1 92 1. Arctonyx annaeus Thomas, Ann. Mag. N.H. 7; 524. Nhatrang, Annam, 

Arctonyx collaris consul Pocock, 1940 

1940. Arctonyx collaris consul Pocock, J. Bombay N.H. Soc. 41: 465. Thaundaung, 
near Toungoo, 4,500 ft.. Lower Burma. Range: Northern Tenasserim to 


Subfamily L u t r i n a e 

Genus LUTRA Brisson, 1762 

1762. Lutra Brisson, Regn. Anim. 13. Alustela Intra Linnaeus. Hopwood (1947, 
P.Z.S. 533-536) would disregard Brisson and date Lutra from Briinnich, 
1 77 1. Zool. Fundamenta, 34, 42, type Mustela lutra Linnaeus. 

1806. Lutris Dumeril, Zool. Analytique, 12. Modification oi Lutra. 

1 81 5. Lutrix Rafinesque, Anal, de la Nature, 59. Substitute for Lutra. 

1865. Barangia Gray, P.Z.S. 123. Lutra sumatrana Gray. 

1865. Lutrogale Gray, P.Z.S. 127. 'The species identified by Gray as monticola Hodg- 
son, which is perspicillata Geoffrey, not monticola Hodgson." Valid as a 

1867. Lutronectes Gray, P.Z.S. 180. Lutronectes whiteleyi Gray = Mustela lutra 

192 1. Hydrictis Pocock, P.Z.S. 543. Lutra maculicollis Lichtenstein, from South Africa. 
Valid as a subgenus. 

3 species in the area covered by this list: 
Lutra lutra, page 275 
Lutra perspicillata, page 277 
Lutra sumatrana, page 277 
Of these, L. sumatrana is nearly extralimital, only touching the region now under 
discussion in Indo-China. L. perspicillata belongs to the genus or subgenus Lutrogale. 
Pocock gave this generic rank, but there seems to be too much tendency to genus- 
splitting in the subfamily, and we provisionally regard it as a subgenus. For characters, 
see Pocock (1941), in which the three species are discussed. 

Subgenus LUTRA Brisson, 1762 

Lutra lutra Linnaeus, 1758 Common Otter 

Appro.ximate distribution of species: widely distributed in the Palaearctic region, 
and in the Indo-Malayan region as far as Java. 

(In detail, known from British Isles, Ireland included, France, Holland, Belgium, 
Spain, Italy, Switzerland, Norway, Sweden, Denmark, Germany, Bohemia, Hun- 
gary, Rumania (? other countries in Europe); Poland; in the U.S.S.R., according to 


i'ai.a[-:arc:tk; a\d Indian mammals 1758-1946 

Bobrinskii it is widely distributed but nearly everywhere rare; it fails to occur only in 
the extreme north-east of European Russia, the extreme north of Siberia, Crimea, 
and in a large part of Kazakstan and the lowlands of Central Asia. Chinese Turkestan, 
Tibet; Japan, Formosa; all the larger states of China, Chihli perhaps excepted; 
Hainan. Ceylon, Southern India, Kashmir to Nepal, Assam, Northern Burma; Indo- 
C^hina, has been recorded from Siam. Sumatra and Java. Asia Minor (B.M.), Persia, 
Palestine. Morocco and Algeria.) 

LiTRA n"rRA LUTRA Linuaeus, 1 758 

1758. Mustda lulia Linnaeus, Syst. Nat. loth ed. /.• 45. Upsala, Sweden. 

1777. Lustra vulgaris Erxleben, Syst. Regn. Anim. /.• 448. Renaming o{ Intra. 

i7C)2. Mustela Lutra piscatoria Kerr, Anim. Kingd. 172. Renaming of lutra. 

(?) 1816. Lutra ftuviatilis Leach, Syst. Cat. Spec. Indig. Mamm. & Birds B.AL 6, 
num. null. 

1827. Lutra vulgaris var. marinus Billberg, Synops. Faunae Scandinaviae, 28. Coasts of 
Scandinavia. Not of Erxleben, 1777. 

1834. Lutra nudipt's Melchior, Den Danske Stats og Norges Pattedyr, 50. Coasts of 
Northern Norway. 

1 8 ^54. Lutra roensis Ogilby, P.Z..S. 1 1 1 . Roe Mills, near Newton Lemavaddy, London- 
derry, Ireland. 

1867. Lutronectes whiteleyi Gra.y, P.Z.S. 181. Japan. 

1887. Lutra lutra var. japonica Nehring, S.B. Ges. Nat. Fr. Berlin, No. 3: 22. Re- 
naming of ivhitelevi. 

(?) iq22. Lutra vulgaris var. baicalensis Dybowski, Arch. Tow. Nauk. Lwow, /.• 349, 
noni. nud. Near Lake Baikal. 

(?) 1922. Lutra vulgaris va.T. amurensis Dybowski, loc. cit. Amur, Ussuri regions, nom. nud. 

(?) 1922. Lutra vulgaris var. kamtschatica Dybowski, loc. cit., nom. nud. Kamtchatka. 

(?) 1936. /.((/ra .i7(7V;f;^(77 Goldman, J. Mamm. ly: 164. Pctropavlovsk, Kamtchatka. 

Range: European and .Siberian range of the species, and Japan (including Kuriles, 
Hondo, .Shikciku, Kiushiu). 

Li'TR.'^ LUTR.\ B.\RANG F. Cuvier, 1823 

1823. Lutra lutra harang F. Cuvier, Diet. Sci. Nat. Paris, 2j: 246. Sumatra. Range 
includes Java, also Annam and Siam. 

Lltr.\ lutr.'\ \ F. Cuvier, 1823 

1823. Lutra nair F. C:u\ier, Diet. Sci. Nat. Paris, 2j: 247. Pondicherry, India. 

1837. Lutra indica Gray, Charlesworth's Mag. N.H. /.• 580. Madras. 

ir)20. Lutra lutra cevlomca Pfihle, Arch. Naturg. 8f;, 9: 72. Nuwara Eliya, Ceylon. 

Range: Ceylon and Southern India (known from Coorg, Nilgiri and Palni Hills). 


1837. Lutra chimnsis Gray, Mag. N.H. /.■ 580. Probably neighbourhood of Canton, 

Sijuthern C-hina. 
1897. Lutra sinensis Trouessart, Cat. Mamm. 283. 
(?) 1007. Lutra hanensis Matschie, Wiss. Ergebn. iMKhner Exped. to China, 10, i: 

150. Hsinganfu, Shensi, China. 
Raiit'c: Chin.i, Hainan and Formosa. 



1839. Lutra monticolus Hodgson, J. Asiat. Soc. Bengal, 8: 320. Nepal. Range: Punjab, 
Kumaon, Nepal, Sikkim, Assam. 


1839. Lutra aurobrunneus Hodgson, J. Asiat. Soc. Bengal, 8: 320. Nepal. 
1865. Barangia ? nepalensis Gray, P.Z.S. 124. Nepal. 

Range: Nepal, at high altitudes, and Garhwal. 

Lutra lutra kutab Schinz, 1844 

1844. Lutra kutab Schinz, Syn. Mamm. 354. Kashmir. Range: to Tibet. 

Lutra lutra angustifrons Lataste, 1885 

1885. Lutra angustifrons Lataste, Actes Soc. Linn, de Bordeaux, jg: 168, 237. Algeria. 
A doubtful form; synonym of/,. /. lutra according to Miller (191 2), but 
available for the North African Otter if it proves racially separable. 

1906. Lutra lutra splendida Cabrera, Bol. Real. Soc. Esp. H.N. Madrid, 6: 360. 
Mogador, Morocco. 

Lutra lutra seistanica Birula, 191 2 

igi2. Lutra lutra seistanica Birula, Ann. Mus. Zool. Acad. Sci. St. Petersb. ij: 274. 

River Gilmend, Seistan, Persia. 
1915. Lutra lutra oxiana Birula, Ann. Mus. Zool. Acad. Sci. St. Petersb. ig: xxi. River 

Pjandsh, Pamir Mountains. 

Range: includes Palestine. 

Lutra lutra meridionalis Ognev, 1931 

193 1. Lutra lutra meridionalis Ognev, Mamm. E. Europe, 2: 527. Surroundings of 
Teheran, Northern Persia. 

Lutra sumatrana Gray, 1865 Hairy-nosed Otter 

Approximate distribution of species: Borneo, Banka, Sumatra, Malay States, north 
to Annam in Indo-China. 

Lutra sumatrana Gray, 1865 

1865. Barangia sumatrana Gray, P.Z.S. 123. Sumatra. (Range as above.) 

Subgenus LUTROGALE Gray, 1865 

Lutra perspicillata Geoffroy, 1826 Smooth-coated Indian Otter 

Approximate distribution of species: Sumatra, Malay States, Indo-China, possibly 
Western Yunnan, Burma, Assam, Nepal Terai, Sind, and southwards to Travancore 
in India. 



1826. Lutra perspicillata I. Gcoffroy, Diet. Class. H.N. 9; 519. Sumatra. 

1827. Lutra simung Lesson, Man. Mamm. 156. Sumatra. 

1839. ■^"''■'z taravensis Hodgson, J. Asiat. Soc. Bengal, 8: 319. Nepal Tcrai. 
1865. Lutra macrodus Gray, P.Z.S. 128. Madras (see Pocock, 1941, 294). 
1879. Lutra ellioti Anderson, Zool. Res. Yunnan, 212. Madras, India. 
Range: as in the species, excepting Sind. 


1940. Lutrogale perspicillata sindica Pocock, J. Bombay N.H. Soc. 41: 517. Chak, 
Sukkur district. Western Sind, India. Range: Indus Valley, from Bahawal- 
pur (Northern Rajputana) to Sind. 

Genus AONYX Lesson, 1827 

1827. Aonyx Lesson, Man. Mamm. 157. Aonyx dtialandi Lesson = Lutra capensis 
Schinz, the Large Small-clawed Otter of Tropical and South Africa. 

1832. Amblonyx Rafinesque, Atlantic J. /.• 62. Amblonyx concolor Rafincsque. Valid as 
a subgenus. 

1842. Leptonvx Lesson, Nouv. Tabl. Regn. Anim. Mamm. /.■ 72. Lutra Iffitonxx Hors- 
ficld = Lutra cinerea Illiger. Not of Swainson, 1821. 

1920. Micraoryx]. Allen, J. Mamm. /.• 24. Lutra cinerea Illiger. 

The name Amblonyx is used as a genus by Pocock (1941) and is so listed by Simpson 
( i945)> ^nd G. Allen (1938) treated the species as a genus under the name Micraonyx. 
However, notwithstanding the differences pointed out by J. Allen in 1920 between 
the small Oriental and the large Ethiopian short-clawed otters, we prefer to follow 
Osgood (1932, Field Mus. N.H. J^ool. 18: 193, et seq.) who in a paper on Indo-Chinese 
Mammals lists the Oriental small-clawed Otter as Aonyx cinerea. Chasen (1940) in- 
cludes cinerea in the genus Lutra, but the short claws of this and allied species are, in 
our opinion, of generic value. 
I species in Asia: 

Aonyx cinerea, page 278 

Subgenus AMBLONYX Rafinesque, 1832 

Aonyx cinerea Illiger, 181 5 Oriental Small-clawed Otter 

.Approximate distribution of species: Southern China (Yunnan, Hainan, Fukien) ; 
Northern Burma, Assam, Sikkim, Nepal, Eastern Punjab; Nilgiri Hills and Coorg, in 
Peninsular India; Indo-China, Malay States, Sumatra, Java, Borneo, Palawan. 

Aonyx cinerea cinerea Illiger, 181 5 

1815. Lutra cinerea Illiger, Abh. .Akad. Phys. Klasse W'iss. Berlin, 1804-11: 90, 99. 

Batavia, Java. 
1823. Lutra leptonyx Horsfield, Zool. Res. Java, pt. 7, pi. Java. 

This race is probably extralimital, although used by both G. Allen for China and 
Osgood for Indri-China. Perhaps their specimens represented the ne.xt race. 



AoNYx ciNEREA coNCOLOR Rafinesque, 1832 

1832. Amblonyx concolor Rafinesque, Atlantic J. /; 62. Garo Hills, Assam. 

1839. Lutra indigitatus Hodgson, J. Asiat. Soc. Bengal, 8: 320. Nepal. 

1855. Aonyx sikimensis Horsfield (Hodgson MS.), Ann. Mag. N.H. 16: 109. Sikkim. 

(?) 1867. Lutra (Hydrogale) swinhoei Gray, P.Z.S. 182. Gawkang Island, near Amoy, 

Southern China. See Pocock (1941, 307, footnote) on status and locality. 
(?) 1920. Amblonyx cinerea fulvus Pohle, Arch. Nat. 85, 9: 133. Lao Key, Tonkin, 

Range: Himalayas to Annam ? and Southern China, west to Kulu f Eastern Punjab). 


1940. Amblonyx cinerea nirnai Pocock, J. Bombay N.H. Soc. 6i: 515. Virajpet, 
Southern Coorg, 3,000 ft., India. Range: Southern India. 

Genus ENHYDRA Fleming, 1822 

1822. Enhydra Fleming, Philos. of Zool. s: iSj. Muslela lutris Linnaeus. 
1816. Pusa Oken, Lehrb. Nat. j, 2: 985. Not of Scopoli, 1777. 
1827. Latax Gloger, Nova Acta Phys. Med. Acad. Caes. Leop. Carol, /j, 2: 51 1. To 
replace Enhydra on the grounds that it was preoccupied by Enhydris Merrem, 
1829. Enydris Fischer, Syn. Mamm. 228. Emendation oi Enhydra Fleming. 
I species: Enhydra lutris, page 279 

Enhydra lutris Linnaeus, 1758 Sea Otter 

Approximate distribution of species : coasts of North-western North America and 
North-Eastern Asia. Southern Kamtchatka (where rare) and Commander Islands 
are the sole U.S.S.R. localities quoted by Bobrinskii. Kurile Islands. 

Enhydra lutris lutris Linnaeus, 1758 

1758. Mustela lutris Linnaeus, Syst. Nat. loth ed. /.• 45, Kamtchatka. 

1777. Lutra marina Erxleben, Syst. Regn. Anim. 445, Kamtchatka. 

(?) 1800. Lutra gracilis Bechstein, Uebers. vierf. Thiere, 2: 408. "Statenland" (accord- 
ing to Hollister, 1921, J. Mammal. 2: 177, the southernmost island of the 
Kurile group is meant). 

1816. Pusa orientalis Oken, Lehrb. Nat. j, 2: 986. 

1827. Lutra stelleri Lesson, Man. Mamm. 156, Kamtchatka. 

1922. Enhydra lutris kamtschatica Dybowski, Arch. Tow. Nauk. Lwow, /.■ 350, nom. mid. 


Genera: Arctictis, page 290 Lchneumia, page 298 

Arctogalidia, page 290 Paguma, page 288 

Chrotogale, page 292 Paradoxurus, page 285 

Cynogale, page 292 Prionodon, page 284 

Genetta, page 283 Viverra, page 280 

Hemigalus, page 291 Viverricula, page 282 
Herpestes, page 292 



This family was divided into two by Pocock, Viverridae and Herpestidae, and the 
former subdivided into numerous subfamilies. So far as the present region is con- 
cerned, Simpson (1945) lists four subfamilies, here retained, with genera as follows: 

Subfamily \"iverrinae 

Tribe \'iverrini Genetta, Viverricula, Viverra. 

Tribe Prionudontini Prionodo7i. 

Subfamily Paradoxurin'ae 

Tribe Arctogalidiini Arciogahdia. 

Tribe Paradoxurini Paradoxurus, Paguma, Arctic/is. 

Subfamily Hemigalinae 

Tribe Hemigalini Himigaliis, Chrotogale. 

Tribe Cynogalini Cynognle. 

Subfamily Herpestinae 

Herpatei, Ichneumia. 

For the characters of the above genera see Pocock (1941). For the Indian Civets, 
see Pocock, 1939, Fauna of British India, i: 331, and for the Indian Mongooses, 1941, 
2: 2. For non-Indian genera see Pocock, 1933, Rarer genera of Oriental Viverridae, 
P.^.S. 969, in which the characters of' Chrotogale and Cvnogale are given; also Pocock, 
1919, Classification of the Mongooses, Ann. Mag. N.H. j; 516 (Herpestes, Ichneumia), 
and Pocock, 1915, P-Z-'^- ^S') where the external characters of Genetta are compared 
with those of its immediate allies. In the igig paper, Mungos is used for forms now 
called Herpestes. A noticeable feature of the skulls of Genetfa in the Palaearctic region 
compared with Viverra (Indian species) and Viverricula is that the last two have the 
sagittal crest strongly developed, whereas in Genetta it is normally weak. Chrotogale, 
with its peculiar incisors and widely open palatal foramina, seems very distinct from 
its nearest ally Hernigalus. Three of the thirteen genera listed above only just come 
into the region now under discussion: Cvnogale and Chrotogale in Indo-China, and 
Ichneumia in Southern Arabia. 

Subfamily V i \- e r r i n a e 

Genus VrVERRA Linnaeus, 1758 

1758. Viverra Linnaeus, Syst. Nat. loth ed. /.• 43. ]'iverra zihetha Linnaeus. 
1933. Moschothera Pocock, J. Bombay N.H. Soc. j6'.- 441. Viverra civettina Blyth. Valid 
as a subgenus. 

2 species in the area covered by this list: 

Viverra megaspila, page 281 
Viverra zihetha, page 281 

Pocock proposed Moichothera as a full genus, but we regard it as being of only sub- 
generic status. Simpson (1945) does not mention it. According to Pocock, Robinson 



and Kloss regarded civettina as a geographical race of megaspila, and we concur with 
that view. For a comparison of the two species here admitted, see Pocock (1939, 344)- 
A third species, V. tangalunga Gray, 1832, which is nea.r zibetha but smaller in size, 
occurs in the Malay States and Islands. 

Subgenus VIVERRA Linnaeus, 1758 

Viverra zibetha Linnaeus, 1758 Large Indian Civet 

Approximate distribution of species: Southern China, from Fukien westwards to 
Yunnan, thence northwards to Szechuan and Southern Shensi; Hainan; Burma, 
westwards through Assam to Nepal; Indo-China, Siam, Malay States. 

Viverra zibetha zibetha Linnaeus, 1758 

1758. Viverra zibetha Linnaeus, Syst. Nat. loth ed. /.• 44. Bengal. 

1830. Viverra undulata Gray, Spic. Zool., pt. 2, 9, pi. 8. Nepal. 

1842. Viverra orientalis or melanurus Hodgson, Calcutta J.N. H. 2: 47. Nepal. 

1842. {Viverra) civettoides Hodgson, loc. cit. 62. 

Range: Nepal, eastwards to South Kamrup in Assam. 

Viverra zibetha ashtoni Swinhoe, 1864 

1864. Viverra ashtoni Swinhoe, P.Z.S. 379. Suykaou, Min Ri\cr, Fukien, Southern 

1907. Viverra Jikhneri Matschie, Wiss. Ergebn. Filchner Exped. to China, /o, i : 192. 

Hinganfu, South-Eastern Shensi, China. 
Range: Chinese range of the species, as given above. 

Viverra zibetha picta Wroughton, 19 15 

1915. Viverra zibetha picta \Vroughton, J. Bombay N.H. Soc. 24: 64. H'Kamti, 500 ft.. 
Upper Chindwin, Northern Burma. 

(?) 1927. Viverra zibetha surdaster Thomas, P.Z.S. 46. Xieng Khouang, Laos, Indo- 

Range: Assam, Northern Burma, Indo-China. 

Viverra zibetha pruinosa Wroughton, 191 7 

1917. Viverra zibetha pruinosa Wroughton, J. Bombay N.H Soc. 24: 64. Thaget, Little 

Tenasserim River, Tenasserim. 
1920. Viverra zibetha sigillata Robinson & Kloss, Rec. Ind. Mus. ig, 4: 176. Bang Nara, 

Patani, Siamese Malaya. 
Range: Tenasserim to Malay Peninsula. 

Subgenus MOSCHOTHERA Pocock, 1933 

Viverra megaspila Blyth, 1862 Large-spotted Civet 

Approximate distribution of species : Burma, Indo-China, Siam, Malay States, 
Western Ghats and Travancore in Peninsular India. 


1862. Viverra megaspila Blyth, J. Asiat. Soc. Bengal, ji: 331. Prome, Lower Burma. 
Range: Burma, Siam, Indo-China, Malav States. 


1862. Viierra civettina Blyth, J. Asiat. Soc. Bengal, j/.- 332. Travancore, Southern 
India. Considered a distinct species by Pocock (1941) and others. 

Genus VIVERRICULA Hodgson, 1838 

1838. ]'ivcrricitla Hodgson, Ann. Mag. N.H. /; 152. Cive.tta xndica GeofFroy 'Viverra 
indica Desmarest). 

I species Viverricula indica, page 282 

Viverricula indica Desmarest, 1817 Rasse, or Small Indian Civet 

Appro,\imate distribution of species: Southern China, from Szechuan eastwards to 
Fukien, also Hainan and Formosa. Ceylon, Peninsular India generally, north to 
Punjab, thence eastwards to Bhutan, Assam, Burma. Indo-China, Siam, Malay 
•States, Sumatra, Java, Bali. ^Introduced in Madagascar and Sokotra.) 

Pocock, 1933, J. Bombav N.H. Soc. j6: 629-631, regarded the name malaccensis 
Gmelin, 1788, Svst. Nat. i: 92, as not valid for the species. As a substitute he 
proposed to use the name indica GeofFroy, 1803, Cat. Mamm. 113. This name is not 
valid from Geoffroy, since, according to Sherborn, Geoffroy's work was never pub- 
lished, and this was admitted by Pocock, 1939, Fauna of British India, Mamm. i: 364 
(footnote), in which it was stated that Desmarest may be regarded as the author of 
the name. But Chasen, 1935, J. Siam Soc. N.H. Suppl. 10: 41, thought the name 
malaccensis should be retained. 

Viverricula i.ndica i.\'dic.\ Desmarest, 181 7 

181 7. Viverra indica Desmarest, Nouv. Diet. X.H. j: 170. India. Range: Southern 
Peninsular India. 

Viverricula indica be.xgale.msis Gray & Hardwickc, 1831 

1831. ]'iverra bengalensis Gray & Hardwicke, 111. Ind. Zool. / : pi. 4. Calcutta, Bengal. 
Range: Calcutta to Gujerat, possibly Sind. 

Viverricula indic.a. pallida Gray, 1831 

1 83 1. I'lverra pallida Gray, Zool. Misc. /.• 17. Probably near Canton, Kwantung, 

Southern China. 
1907. Viverricula hanensis Matschie, Wiss. Ergebn. Filchncr Exped. to China, /o, i : 

196. Hankow. Southern China. 
191 1. Viverricula pallida taivana Schwarz, Ann. Mag. N.H. j: 637. Formosa. 
Range: Szechuan, Yunnan, Fukien, etc., in Southern China; and Formosa. 



1898. Viverricula malaccensis deserti Bonhote, Ann. Mag. N.H. /.• 120. Sambhar, Raj- 
putana, India. 

Viverricula indica thai Kloss, 19 19 

1919. Viverricula malaccensis thai Kloss, J. N.H. Soc. Siam, j.- 352. Prapatom, west of 
Bangkok, Siam. Range: Burma, Siam, Indo-China; possibly the form listed 
as V. malaccensis malaccensis from Hainan in G. Allen, 1938, Mamm. China & 

Viverricula indica mayori Pocock, 1933 

1933. Viverricula indica mayori Pocock, J. Bombay, N.H. Soc. 56'.' 632. Maha Oya, 
Eastern Province, Ceylon. 

Viverricula indica wellsi Pocock, 1933 

1933. Viverricula indica wellsi Pocock, J. Bombay N.H. Soc. ;^6: 640. Kangra, 2,000 ft., 
Punjab, Northern India. Range: Kangra to Kumaon. 

Viverricula indica baptistae Pocock, 1933 

1933. Viverricula indica baptistae Pocock, J. Bombay N.H. Soc. 56'.- 643. Hasimara, 
Bhutan Duars, India. Range: to Assam. 

Genus GENETTA Oken, 18 16 

1816. Genetta Oken, Lehrb. Nat. j, 2: loio. Viverra genetta Linnaeus (see page 3). 
1816. Genetta Cuvier, Regn. Anim. /: 156. Viverra genetta Linnaeus. 
1841. Odmaelurus Gloger, Gemeinn. Hand. u. Hilfsbuch der Nat. /.• 72. Viverra genetta 

I species in the area 'covered by this list: 
Genetta genetta, page 283 

This genus, several species of which occur in Ethiopian Africa, was revised by 
Schwarz, 1930, Rev. ^ool. Bot. Afr. ig, 2 : 276-286. Only one species enters the present 

Genetta genetta Linnaeus, 1758 European Genet 

Approximate distribution of species: France, Spain, Balearic Islands; also has 
been recorded from Germany, Switzerland and Belgium. Palestine, Arabia. Morocco, 
Algeria, Libya, Africa south of the Sahara, southwards to the Transvaal and at least 
to Clanwilliam in West Cape Province; east to Somaliland, and west to Senegal and 

Genetta genetta genetta Linnaeus, 1758 
1758. Viverra genetta Linnaeus, Syst. Nat. loth ed. /.■ 45. Spain. 
1816. Viverra Genetta hispanica Oken, Lehrb. der Nat. j, 2: 10 10. Ronda, Malaga, 

T 283 

rALAEARCniC: and IXDIAX mammals 1758-1946 

Genetta cenetta genetta [conld.] 

1816. Viverm Genetta gallica Okcn, loc. cit. loin, alternative name for hispanica, n(it of 

Kerr, 1792. 
1827. Gnietta vulgaris Lesson, Man. Mamni. 17:5. Renaminc; of ^fncZ/a. 
(?) 1830. Genetta enmmnni.s Burnett, Quart. J. Sci. Lit. Art. iSsg, 2: 349, nom. mid. 
1897. Genet/a melas Graelis, Mem. ReaL Acad. Sci. NLadrid, ly: 175. Sierra Morena, 

(?) 1905. Genetta /leinnstilae Cabrera, BoL ReaL See. Esp. H.N. 266. El Pardo, near 

Madrid, Spain. 
Range: Spain. 

Ge,nett.\ genetta .xfra F. Cuvier, 1825 

1825. Genetta afia F. C^uvier, in Cuvier & Gcoflroy, H.N. ^L^mm. pt. 52, pL 195; and 

pt. 51, text. Barbary. 
1842. Genetta genetta barbara H. Smith, Jardinc's Nat. Library, ALamm. 25- ^7^- 

1857. Genetta bonaparli Loche, Rev. Mag. Zooi. 9, 2: 385, pi. 13. Algeria. 
Range : \\ cstcrn \Iorocco, Algeria, Tunis, Libva. 

Genetta genetta bale.^rica Thomas, 1902 

1902. Genetta genetta halearica Thomas, Ann. ALag. N.H. 10: 162. Inca, Majorca, 
Balearic Islands. 

Genett..\ genetta rhodaxica Matschie, 1902 

1902. Genetta rhodamca .\Latschie, Verhandl. 5th Int. Zool. Congr. Berlin, 11 39. 
Montpellicr, Herault, France. Range: South-Western France. 

Genetta genetta granti Thomas, 1902 

1902. Genetta grantii Thomas, Ann. Mag. N.H. 10: 487. Azraki Ravine, Haushabi, 
5,200 ft., Arabia. 

Genetta genetta terraesanctae Neumann, 1902 

ig02. Genetta terraesanelae Neumann, S.B. Ges. Nat. Fr. Berlin, 183. Mt. Carmel, 


Genus PRIONODON Horsfield, 1822 

1822. Prionodnn Horsfield, Zool. Res. Java, pt. 3. Felts graeilis Horsfield (= P. linsang 

graeilis, from Java). 
1839. Linsang Muller, Verb. Nat. Ges. Nederl. /, Taf (3) : 28. Felis gracilis Horsfield. 
1842. Pnodovtes Lesson, Nou\-. Tabl. R. Anim. bo. Felts gracilis Horsfield. Not of 

Cuvier, 1827. 
1896. Linsanga Lydekkcr, Geogr. Hist. Mamm. 20. Emendation of Linsang. 
1925. Pardictis Thomas, P.Z.S. 498. Prionodon pardicolor Hodgson. Valid as a subgenus. 

2 species: Prionodon linsang, page 283 

Prinitodon pardicolor, page 28", 
Pocock I 1930. 33*1) gi\cs a key to the species. He ignores Pardictis whkh Simpson 
(19451 lists as a lull genus, Osgood (1932) as a subgenus. \Ve propose to follow 
Osgood . 



Subgenus PRIONODON Horsfield, 1822 

Prionodon linsang Hardwicke, 182 1 Banded Linsang 

Approximate distribution of species: Tenasserim, Malay States, Sumatra, Java, 

Prionodon linsang linsang Hardwicke, 182 1 

1 82 1. Vivenal linsang Hardwicke, Trans. Linn. Soc. London, 75; 236, pi. 24. 

1878. Prionodon maculosus Blanford, Proc. As. Soc. Bengal, 71. Bankachon, Southern 


Range: Tenasserim to Sumatra. 

Subgenus PARDICTIS Thomas, 1925 

Prionodon pardicolor Hodgson, 1841 Spotted Linsang 

Approximate distribution of species: Nepal, Assam, Northern Burma, Indo- 

Prionodon pardicolor pardicolor Hodgson, 1841 

1841. Prionodon pardicolor (sic) Hodgson, Calcutta J.N. H. 2: 57. Nepal. 
1844. Viverra perdicator Schinz, Syn. Mamm. /.• 366. Error ior pardicolor. 
1863. Prionodon pardochrous Gray, Cat. Hodgsons Coll. B.M. 4, nom. nud. 

Ranges to Assam and Northern Burma. 

Prionodon pardicolor presin.a Thomas, 1925 

1925. Pardictis pardicolor presina Thomas, P.Z.S. 499. Ngai-tio, 4,800 ft. .Tonkin, Indo- 
China. Osgood thought this was a synonym of the typical race. 

Subfamily Paradoxurinae 
(as understood by Simpson, 1945) 

Genus PARADOXURUS Cuvier, 1821 

1 82 1. Paradoxurus Cuvier, in Cuvier & Geoffroy, H.N. Mamm. 2, 24: Martre des 

Palmiers, 5. Paradoxurus typus Cuvier = Viverra hermaphrodila Pallas. 
1835. Platyschista Otto, Nov. Act. Acad. Caes. Leop. Carol, ij: 1089. Platyschista 

pallasii Otto = Viverra hermaphrodila Pallas. 
1864. Bondar Gray, P.Z.S. 531. Viverra hondar Desmarest. 

1864. Macrodus Gray , P.Z.S. 536. Paradoxurus macrodus Gray = Viverra musangajavanica 
Horsfield (the Javan race o^ hermaphroditus) . 



3 species: Paradoxurus hermaphroditus, page i86 
Paradoxurus jerdoni, page 288 
Paradoxurus zeylo7iensis, page 288 

Pocock retains three species as above, and compares them (1939, 380). P. jerdoni 
seems very close to zn'lonensis, and might well be considered as a subspecies of it. 

Paradoxurus hermaphroditus Pallas, 1 777 Common Palm Civet, or Toddy Cat 
Approximate distribution of species: Hainan and Kwantung, in Southern China; 
Burma and Assam westwards to Kashmir, thence southwards through Peninsular 
India to Ceylon; Indo-China, Siam, Malay States, Sumatra, many small adjacent 
islands, Java, Borneo, to Celebes, the Philippines, Timor, Ceram and the Kei 
Islands (perhaps introduced in some of the easternmost islands just quoted). 

Paradoxurus hermaphroditus hermaphroditus Pallas, in Schreber, 1777 
1777. Mvcrra hermaphrodita Pallas, in Schreber, Saugeth. 3: 426. ? India. 

1820. Viverra nigra Desmarest, Mamm. 208. (Not of Peale & Beauvois, 1796.) Pondi- 

cherry, India. 

1821. Paradoxurus lypus F. Cuvier & Geoffroy, H.N. Mamm. pt. 24, 5. Pondichcrry. 
1832. Paradoxurus tvpus var. fuliginosus Gray, P.Z.S. 65. Southern India. 

1841. Paradoxurusjelinus Wagner, Schrcb. Saugeth. Suppl. 2: 349. India. (Composite: 

composed partly oi hermaphroditus and partly of pallasi.) 
1885. Paradoxurus niger Blanford, P.Z.S. 792. Pondichcrry, India. 
Range: Ceylon and Southern India, as far north as the Narbada River. 

Paradoxurus hermaphroditus bondar Desmarest, 1820 

1820. Viverra hondar Desmarest, Mamm. 210. Bengal. 

1832. Paradoxurus pennantii Gray, P.Z.S. 66. Higher Province of Bengal. 

1832. Paradoxurus crossi Gray, P.Z.S. 67. India. 

1836. Paradoxurus hirsutus Hodgson, Asiat. Res. ig: 72. Nepal Terai. 

1855. Paradoxurus stric/us Horsfield (Hodgson MS.), Ann. Mag. N.H. j6: 105. Nepal 

Range: Kumaon, Nepal Terai and district. 

Paradoxurus hermaphroditus pallasi Gray, 1832 

1832. Paradoxurus pallasii Gray, P.Z.S. 67. India. 

1820. Viverra prehensilis Desmarest, Mamm. 208, not of Kerr, 1792. Bengal. 

1855. Paradoxurus quadriscriptus Horsfield (Hodgson MS.), Ann. Mag. N.H. 16: 106. 

Nepal (Hills). 
1864. Paradoxurus nigrijrons Gray, P.Z.S. 535. India. 

1910. Paradoxurus vicinus Schwarz, Ann. Mag. N.H. 6: 230. Probably Assam. 
Range: Nepal, Sikkim, Assam, Upper Burma. 

Paradoxurus hermaphroditus nictitatans Taylor, 1891 

1891. Paradoxurus nictitatans Taylor, J. Bombay N.H. Soc. 6: 429, pi. Kondmals, 

Orissa Division of Southern Bengal. 
(?) 1829. Paradoxurus leucopus Ogilby, Zool. J. 4: 301. "Probably some part of the 

East Indies." 



Paradoxurus hermaphroditus minor Bonhote, 1903 

1903. Paradoxurus minor Bonhote, Fasc. Malay Zool. /; 9. Kampong Jalor, Lower 
Siam. According to Pocock, occurs in Tenasserim. For status of this form 
see Chasen, 1940, Handlist Malaysian Mamm. 95, 96. 

Paradoxurus hermaphroditus cochinensis Schwarz, 1911 

191 1. Paradoxurus cochinensis Schwarz, Ann. Mag. N.H. j: 635. Saigon, Cochin- 

Paradoxurus hermaphroditus exitus Schwarz, igii 

191 1 Paradoxurus exitus Schwarz, Ann. Mag. N.H. y: 636. Fumai, east of Canton, 
Kwantung, Southern China. 

Paradoxurus hermaphroditus senex Miller, 191 3 

1913. Paradoxurus hermaphroditus senex Miller, Smiths. Misc. Coll. 61, 21: 3. Domel 
Island, Mergui Archipelago. 

Paradoxurus hermaphroditus fallens Miller, 1913 

1913. Paradoxurus hermaphroditus palleru Miller, Smiths. Misc. Coll. 61, 21: 4. Kisser- 
aing Island, Mergui Archipelago. 

Paradoxurus hermaphroditus pugnax Miller, 19 13 

1 91 3. Paradoxurus hermaphroditus pugnax Miller, Smiths. Misc. Coll. 61,21: 4. Sullivan 
Island, Mergui Archipelago. 

Paradoxurus hermaphroditus sacer Miller, 1913 

191 3. Paradoxurus hermaphroditus sacer Miller, Smiths. Misc. Coll. 61, 21: 4. St. 
Matthew Island, Mergui Archipelago. 

Paradoxurus hermaphroditus pulcher Miller, 19 13 

1913. Paradoxurus hermaphroditus pulcher Miller, Smiths. Misc. Coll. 61, 21: 5. Clara 
Island, Mergui Archipelago. 

Paradoxurus hermaphroditus laotum Gyldenstolpe, 191 7 

19 1 7. Paradoxurus hermaphroditus laotum Gyldenstolpe, K. Svenska. Vet. Akad. Handl. 

57, 2 : 26. Chieng Hai, North-Western Siam. 
191 7. Paradoxurus birmanicus Wroughton, J. Bombay N.H. Soc. 25.- 51. Mingun, near 

Sagaing, Upper Burma. 

Range: Burma (Mandalay and Chindwin to Tenasserim), Siam, Indo-China and 

Paradoxurus hermaphroditus scindiae Pocock, 1934 

1934. Paradoxurus hermaphroditus scindiae Pocock, J. Bombay N.H. Soc. ^7: 176. Guna, 
in Gwalior (about 40 miles north oflatitude 24^), India. 



Paraddxi'ris hermaphrodites laneus Pocock, 1934 

11)14. PaiadoMiius htnnaphwditus laneus Pocock, J. Bombay N.H. Soc. ^y: 178, fig. 4b. 
Gopalpur, 5,200 ft., Kangra, Punjab. 

Paradoxurus hermaphroditus vellerosus Pocock, 1934 

1034. Panidnxu)u.\ lurmaphnHlitin relkroitis Pocock, J. Bombay N.H. Soc. 37; 181. 

Paradoxirv-S hermaphroditus milleri rinm. nov. 

1013. Paradoxurus hermaphroditus fuscus Miller, Smiths. Misc. Coll. 6'/, 21: 3. James 
Island, Mergui Archipelago. Not/ia«« Kclaart, 1852. 

Paradoxurus zeylonensis Pallas, in Schreber, 1778 Golden Palm Civet 

.Approximate distribution of species: Ceylon. 

Paradoxurus zeylonensis Pallas, in Schrcljcr, 1778 

1778. Vmria rexhnensis Pallas, in Sclin-bcr, Saugcth. ;,': 451. C:cvlon. 

1788. Viriira .yvlanica Gmelin, Syst. Nat. 13th cd. /.■ 89. Ceylon. 

,?) 1822. Paradoxurus aureus F. Cuvier, Mem. Mus. H.N. Paris, c/: 48, pi. 4. Locality 

1852. Paradoxurus rjylariirus with Juseus or monlamis Kelaart, Prodr. Faun. Zeylan. 

39-40. Newera Eliya, Cleylon. 

Paradoxurus jerdoni Blanford, 1885 Jerdon's Palm Civet 

Approximate distribution of species: Palni Hills, .\ilgiri Hills, Coorg and Tra\an- 
core in Southern India. 

Paradoxurus jerdoni jerdom Blanford, 1885 

1885. Paradoxurus jerdom Blanford, P.Z.S. (J13, 802. Kod.iikanal, I'alni Hills, Southern 

Paradoxurus jerdoni caniscus Pocock, 1933 

1933. Paradoxurus jerdoni eaniseus Pocock, J. Bomba\ N.H. Soc. j-O; 865. \ irajpet, 
3,000 ft.. Southern Coorg, India. 

Genus PAGUMA Gray, 1831 

1831. Paguma Grav, P.Z.S. i8jo-ji: or,, (uilo larralus Hamilton-Smith. 
1837. Amliliodon jourddn. C.R. Acad. Sci. Paris, j: 445. Paradoxurus jourdami Gray 
' -- the Malac I .ui race oi Pnt^uma larvata). 

I species: Pat^uina lairala, page 289 
Pocock I I03r), 41b) also lists a species /'. lariigera Hodgson, based on an "imperfect, 
no doubt miniature" skin without skull iroiii the "northern region ol", sub- 
seciucntU said to be from Tingrce, Tibet. If its skull is not known, presumably its 
gcncrir positii.n is not certainly known, as Paouma differs from Paradoxurus chiefly in a 
cranial ( haracter (the Iciiglli of the palate). \Ve propose to regard it as ineertae sedis. 



Paguma larvata Hamilton-Smith, 1827 Masked Palm Civet 

Approximate distribution of species: China, from Fukien and south-east coast 
westwards to Yunnan, thence northwards to Szechuan, Southern Shensi and Chihli 
(Pekin); Hainan, Formosa. Burma and Assam westwards to Kashmir; Andaman 
Islands. Indo-China, Siam, Malay States, Sumatra, Borneo. 

Paguma larvata larvata Hamilton-Smith, 1827 

1827. Gulo larvatus Hamilton-Smith, Griffith's Cuvier Anim. Kingd. 2: 281, pi. 

Locality unknown. 
1907. Paguma reevesi Matschie, \Viss. Ergebn. Exped. Filchner to China, 10, i: 183. 

Hing-an-fu, China. 
1 92 1. Paguma larvata nra/w Thomas, Ann. Mag. X.H. 8: 618. Ichang, Hupeh, China., 
Range: eastern parts of Southern China, west to Szechuan. 

Paguma larvata gray: Bennett, 1835 

1835. Paradoxurus graji Bennett, P.Z.S. 118. India. 

1836. Paradoxurus nipalensis Hodgson, Asiat. Res. ig: 76. Nepal. 
Range: Nepal, west to Kumaon and Garwhal. 

Paguma larvata taivana Swinhoe, 1862 

1862. Paguma larvata var. taivana Swinhoe, P.Z.S. 354. Formosa. Range includes 
Botel Tobago. 

Paguma larvata tytleri Tytler, 1864 

1864. Paradoxurus tytlerii Tytler, J. Asiat. Soc. Bengal, 5^.- 188. Viper Island, Port 
Blair, South Andaman Island. 

Paguma larvata robusta Miller, 1906 

1906. Paradoxurus robustus Miller, Proc. Biol. Soc. Washington, ig: 26. Trang, Lower 
Siam. Ranges to Tenasserim. 

Paguma larvata hainana Thomas, 1909 

1909. Paguma larvata hainana Thomas, Ann. Mag N.H. 5.- 377. Five Finger Moun- 

tains (~\Vuchih), Island of Hainan, Southern China. 

Paguma larvata intrudens Wroughton, 19 10 

1910. Paguma larvata intrudens \Vroughton, J. Bombay N.H. Soc. ig: 793. Sima, 

Myitkyina, Upper Burma. 

1919. Paguma larvata vagans Kloss, J.N.H. Soc. Siam, jj.- 73. Sikawtur, 40 miles north- 
west of Raheng, 1,500 ft., Siam. 

1 92 1 . Paguma larvata yunalis Thomas, Ann. Mag. N.H. 8: 617. Yenyuensien, Southern 
Szechuan, China. 

Range: Szechuan, Yunnan, Northern Burma to Shan States, Siam; Laos, Annam 
and Tonkin, in Indo-China. 

Paguma larvata wroughtoni Schwarz, 19 13 

1913. Paguma grayi wroughtoni Schwarz, Ann. Mag. N.H. 12: 289. Gharial, near 
Murree, Northern Punjab, India. Range: Kumaon to Kashmir. 



Paguma larvata janetta Thomas, 1928 

1928. Paguma leucomystax janetta Thomas, Ann. Mag. N.H. 2: loi. Bankachon, 
Southern Tenasserim. 

Paguma larvata neglecta Pocock, 1934 

1934. Paguma larvata neglecta Pocock, J. Bombay N.H. Soc. 27- 334- Mokokchung, 

4,500 ft., Naga Hills, Assam. Range: low-lying districts of Nepal, Sikkim, 

Assam, Chin Hills and Arakan, Western Burma. 


1939. Paguma larvata nigriceps Pocock, Fauna Brit. India, Mamm. /.• 424. Nam 
Tamai, Upper Burma. 

{Incertae sedis: see remarks above) 

Paguma (?) lanigera Hodgson, 1836 
1836. Paradoxurus lanigerus Hodgson, Asiat. Res. ig: yg. 

1841. Paradoxurus laniger Hodgson, J. Asiat. Soc. Bengal, 10: 909. "Northern region 
of Nepal," subsequently said to be from Tingree, Tibet. 

Genus ARCTICTIS Temminck, 1824 

1824. Arctictii Temminck, Mon. Mamm. /, Tabl. Method, xxi. Viverra binturong 

1824. Ictides F. Cuvier, Dents Mamm. 252. Viverra binturong Raffles. 

I species: Arctictis binturong, page 290 

Arctictis binturong Raffles, 1821 Binturong 

Approximate distribution of species: Burma (possibly Assam, Bhutan, Nepal, 
Sikkim); Indo-China, Siam, Malay States, Sumatra, Java, Borneo, Palawan. 

Arctictis binturong binturong Raffles, 1821 

182 1. Viverra'^ binturong Raffles, Trans. Linn. Soc. London, /j.' 253. Malacca. 
1916. Arctictis gairdnen Thomas, Ann. Mag. N.H. ij: 270. Sai Yoke, South-Western 

Ranges to Tenasserim. 

Arctictis binturong albifrons F. Cuvier, 1822 

1822. Paradoxurus albifrons F. Cuvier, Mem. Mus. H.N. Paris, g: 44, 48. Bhutan, 

Eastern Himalayas. Range: Upper Burma, Indo-China. 

Genus ARCTOGALIDIA Merriam, 1897 

1864. Arctogale Gray, P.Z.S. 542. Not Arctogale Kaup, 1829. Paradoxurus trtvirgatus 

1897. Arctogalidia Merriam, Science, j.' 302. New name for Arctogale Gray, pre- 
occupied. Paradoxurus trivirgatus Gray. 

I species: Arctogalidia trivirgata, page 291 


Arctogalidia trivirgata Gray, 1832 Small-toothed Palm Civet 

Approximate distribution of species: Assam, Burma, Indo-China, Siam, Malaya, 
Sumatra, and numerous small adjacent islands, Java, Borneo. 

(Arctogalidia trivirgata trivirgata Gray, 1832. Extralimital) 
1832. ParadoxuTus trivirgatus Gray, P.Z.S. 68. Buitenzorg, Western Java. 

Arctogalidia trivirgata leucotis Horsfield, 1851 

1 85 1. Paradoxurus leucotis Horsfield, Cat. Mamm. E. India Co. 66. Tenasserim. 
1877. Paradoxurus preheruilis Sclater, P.Z.S. 681, pi. 71. Not of Desmarest, 1820. 
Range: Burma, Siam, Tenasserim, Kings Island, Mergui Archipelago. 

Arctogalidia trivirgata major Miller, 1906 

1906. Arctogalidia major Miller, Proc. Biol. Soc Washington, ig: 25. Trang, Lower 
Siam. Occurs Laos and Tonkin, in Indo-China, according to Tate. 

Arctogalidia trivirgata macra Miller, 19 13 

1913. Arctogalidia macra Miller, Smiths. Misc. Coll. 61: 6. Domel Island, Mergui 

Arctogalidia trivirgata millsi Wroughton, 1921 

192 1. Arctogalidia millsi Wroughton, J. Bombay N.H. Soc. 2y: 600. Mokokchung, 
Naga Hills, 5,000 ft., Assam. 

Subfamily Hemigalinae 
(As understood by Simpson, 1945) 

Genus HEMIGALUS Jourdan, 1837 

1837. Hemigalus Jourdan, C.R. Acad. Sci. Paris, 5.- 442. Hemigalus zebra Gray = 
Viverra hardwickii Gray. 

I species in the area covered by this list: 
Hemigalus derbyanus, page 291 

Hemigalus derbyanus Gray, 1837 Banded Palm Civet 

Approximate distribution of species : Tenasserim, Malay Peninsula, Sumatra and 
some of the islands to the west of it, Borneo. 

(Hemigalus derbyanus derbyanus Gray, 1837. Extralimital) 

1837. farflf/oAruraj^rfer^varzMjGray, Charlesworth'sMag. N.H. /.• 579. Malay Peninsula. 
(?) 1837. Paradoxurus? zebra Gray, loc. cit. No locality. 



Hemigalus derbyanus i.ncursor Thomas, 1915 

1 9 15. Hfmigciliis (hrhianus (sic) incursor Thomas, ]. Bombay N.H. Soc. I'jj; 613. 
Bankarhoii, \'icti>ria Piovincc, Tcnasscrim. 

Genus CHROTOGALE Thomas, 1912 
1912. Chrotoiidlf Thomas, Abstr. P.Z.S. 17; P.Z.S. 499. Chrotogale owstoni Thomas. 
I species: Chroloaalc owsloni, page 292 

Chrotogale owstoni Thomas, 191 2 Owston's Banded Civet 

Approximate distribution of species: Tonkin and Laos, in Indo-phina. 

Chrotogale owstom Thomas, 19 12 

1912. Chrotogale owsloni Thomas, Abstr. P.Z.S. 17; P.Z.S. 500. Yen Bai, Songkoi 
River, Tonkin, IndoChina. 

Genus CYNOGALE Gray, 1837 

1837. Cvnogale Crux, P.Z.S. i8;]6: 88. Mag. X.H. /, i8;^y: ^jct.Cynogale bennettiiGrdiy. 

1838. Potamophilus Miiller, Tijdschr Nat. Gesch. Phys. 5.- 140. Potamophilus barbatus 

Miiller = C'vnogale bennettii Gray. 

I species: Cytwgale bennetti, page 292 

Cynogale bennetti Gray, 1837 Otter-Civet 

.\ppro.\imatc distribution of species: Indo-China, Malay States, Sumatra, Borneo. 

Pocock separated the Indo-Chinese representative as a distinct species, but until 
more specimens come to hand vvc prefer to regard this very imperfectly-known form 
as a subspecies. 

Cyxog.^le be.nnetti bex.xetti Gray, 1837. Extralimital) 
1837. Cvnogali- hauuilii Gray, P.Z.S. i8j6: 88. Sumatra. 

Cynogale BE.\.\ETn lowei Pocock, 1933 

1933. Cynogale lowei Pocock, P.Z.S. 1034, fig. Backan, 500 ft.. Tonkin, Indo-China. 

Subfamily H c r p c s t i n a e 

Genus HERPESTES Illigcr, 181 1 

lyqq. Ichneiinion Laccpede, Tabl. Div. Ord. Gen. Mamm. 7, not of Linnaeus, 1758. 
i(')ii. Heifieiles llliger, Prodr. Syst. Mamm. et Avium, 133, misprint, corrected to 

llerjnsles, 302. I'nerra ichneumon Gmclin. 
1822. Mangiisia Horsfield, Zool. Res. Java, unpaged, pt. 5. Ichneumon javamcus 




1837. Urva Hodgson, J. Asiat. Soc. Bengal, 6: 561. Gulo una Hodgson. 

1841. Mesobema Hodgson, J. Asiat. Soc. Bengal, 10: 910. Gulo urva Hodgson. 

1865. Calogale Gray, P.Z.S. 1864: 560. Herpesles nepalensis Gray = Mangusta auro- 

punctatus Hodgson. 
1865. Caliclis Gray, P.Z.S. 1864: 564. Herpesles smithii Gray. 
1865. Taeniogale Gray, P.Z.S. 1864: 569. Herpesles vilticollis Bennett. 
1865. Onjchogale Gray, P.Z.S. 1864: 570. Cyniclis maccarthiae Gray. 

"Mungos Cuvier & Geoffroy" of some earlier authors, but Mungos Cuvier & 
Geoffrey, 1795, Mag. Encycl. 2: 184, is now restricted to the Banded Mongoose of 
Africa and its immediate allies. 

8 species in the area covered by this list: 

Herpesles auropiinctalus, page 295 
Herpesles edwardsi, page 295 
Herpesles fuscus, page 297 
Herpesles ichneumon, page 294 
Herpesles javanicus, page 294 
Herpesles smilhi, page 296 
Herpesles urva, page 298 
Herpesles vilticollis, page 298 

Pocock (1941) recognized only six species in India, regarding auropunclatus as a race 
oi javanicus, which he said ranged from Persia through Northern India to Java. But 
Chasen, 1940, Handlisl Malaysian Mammals, 103, states: "Two species of this group 
distinguished chiefly by size occur in the Malay Peninsula; only one can be the local 
representative oi javanicus, and it appears to be the larger form. H. auropunctalus is the 
earliest name for the other association." Therefore, auropunclalus is given specific 
status here. Pocock gave measurements (1941, 34) for various extralimital races of 
his javanicus; most of these, and the Indian forms, seem to be auropunclalus; but 
possibly exilis, which was named from Annam, may be taken as rc-preicntmg javanicus 
in the region now under discussion. The remaining species, H. ichneumon, occurs in 
North Africa, Spain and Palestine, and was not dealt with by Pocock in his work on 
mammals of British India. So far as we can see, there are three groups of Herpesles 
Mongooses in the Palaearctic and Indian regions: ichneumon group (large, as judged 
by size of skull, neck not striped, colour grey with black tailtip; chiefly African); 
vitlicollis group, about as large, but neck conspicuously striped, containing the two 
distinct species vilticollis and urva which are compared by Pocock (1941, 7), and the 
edwardsi group, containing five medium or small species (as judged by size of skull), 
three of which occur together in Ceylon, and the characters of which are dealt with 
by Pocock (1941, 7), but it must be added that auropunclalus as here understood and 
following Chasen averages smaller than javanicus as here understood. Three other 
species, o- ly two of which are available for examination, are listed by Chasen (1940) 
from the Malay region. Of these, H. semilorqualus is very close to H. urva, possibly even 
only a race of it, but H. brachyurus is quite distinct, with the tail proportionately 
shorter than is usual in the other species, and with no neckstripes. 


Herpestes ichneumon group 

Herpestes ichneumon Linnaeus, 1758 Egyptian Mongoose, or Ichneumon 

Approximate distribution of species: Spain and Portugal; Palestine; Morocco, 
Algeria, Egypt; Ethiopian Africa, from Kenya and Nigeria south to South- West 
Africa, Trans\aal, Natal and Knysna in Cape Province. 

Herpestes ichneumon ichneumon Linnaeus, 1758 

1758. Viverra ichneumon Linnaeus, Syst. Nat. loth ed. /.■ 43. Egypt "ad ripas Nili". 

1799. Ichneumon pharaon Lacepede, Tabl. Div. Ord. Gen. Mamm. 7. 

1808. Ichneumon aegypliae Tiedemann, Zool. /.' 364. 

1818. Ichneumon major E. Geoffroy, Descript. Egypte, 2: 139 (footnote). Egypt. 

Range: Egypt and Palestine. 

Herpestes ichneumon numidicus F. Cuvier, 1834 

1834. Ichneumon numidicui Cuvier, H.N. Mamm. pt. 68, pi. 191, and text. Algeria. 
Range : Northern Morocco, Algeria. 

Herpestes ich.neumon widdringtoni Gray, 1842 

1842. Herpestes widdringtonii Gray, Ann. Mag. N.H. 9, i : 50. Sierra Morena, Spain. 
1909. Herpestes ichneumon vav.ferruginea Seabra, Bull. Soc Portugaise. Sci. Nat. .'.• 286. 

Alemtejo, Portugal. Not of Blanford, 1874. 
1909. Herpestes ichneumon var. dorsalis Seabra, loc. cit. Ribatejo, Portugal. 
1909. Herpestes ichneumon var. grisea Seabra, loc. cit. Ribatejo, Portugal. Not of 

Geoffroy, 1818. 
1912. Mungos widdringtonii Miller, Cat. Mamm. \V. Europe, 441. 
Range: Spain and Portugal. 

Herpestes ichneumon sangronizi Cabrera, 1924 

1924. Herpestes ichneumon sangronizi Cabrera, Bol. Real. Soc. Esp. H.N. Madrid, 24: 
217. Mogador, Morocco. 

Herpestes edwardsi group 

Herpestes javanicus Geoffroy, 1818 Javan Mongoose 

Approximate distribution of species: ? Indo-China, Siam, Malay States, Java. 

(Herpestes javanicus javanicus Geoffroy, 181 8. Extralimital) 

1818. Ichneumon javanicus E. Geoffroy, Descr. Egypte, 2: 139. Western Java. 

Herpestes (?) javanicus exilis Gervais, 1841 

1 84 1. Herpestes c.xilis Gervais, Voy. Bonite, /.• 32, pi. 3, figs. 7-9. Tourane, Annam, 

(?) 18G1. Herpestes rutilus Gray, P.Z.S. 136. Cambodia, Indo-China. 



Herpestes javanicus peninsulae Schwarz, 1910 

1910. Mungos exilis peninsulae Schwarz, Ann. Mag. N.H. 6: 231. Bangkok, Siam. 

igiy. Mungos incertus Kloss, J. Fed. Malay States Mus. j: 125. Ongut, Trang, Lower 

Range: Siam, Malay Peninsula. 

Herpestes auropunctatus Hodgson, 1836 Small Indian Mongoose 

Approximate distribution of species : ? Northern Arabia, Persia, Iraq, Afghanistan ; 
Kashmir, south to Gujerat, Sind and Orissa, east to Nepal, Assam and Burma; 
Hainan; Siam, Malay States. 

Herpestes auropunctatus auropunctatus Hodgson, 1836 

1836. Mangusta auropunctata Hodgson, J. Asiat. Soc. Bengal, 5.- 235. Nepal. 

1837. Herpestes nepalensis Gray, Charlesw. Mag. N.H. /; 578. Northern India. 
Range: Kashmir to Manipur and Orissa. 

Herpestes auropunctatus pallipes Blyth, 1845 

1845. Mangusta pallipes Blyth, J. Asiat. Soc. Bengal, i ^: 346. Kandahar, Afghanistan. 
1864. Herpestes persicus Gray, P.Z.S. 554. Mohammerah, Western Persia. 
1914. Mungos auropunctatus helvus Ryley, J. Bombay N.H. Soc. 22: 661. Deesa, 
Palanpur, Gujerat, India. 

Range: Iraq and perhaps Northern Arabia, Afgham'stan, Persia, ? Baluchistan, Sind, 
Punjab, Palanpur, in Western India. 

Herpestes auropunctatus birmanicus Thomas, 1886 

1886. Herpestes auropurwtatus birmanicus Thomas, Ann. Mag. N.H. ij: 84. Pegu, 
Burma. Range: Burma, from Toungoo to Tenasserim. 

Herpestes auropunctatus rubrifrons J. Allen, 1909 

1909. Mungos rubrifrons ]. Allen, Bull. Amer. Mus. N.H. 26: 240. Mount Wuchih, 

Island of Hainan, Southern China. 
1 94 1. [H. javanicus) nigrifrons Pocock, Fauna Brit. India, Mamm. 2: 34. (? lapse for 


Range: Hainan and possibly Kwantung, Southern China. 

Herpestes auropunctatus siamensis Kloss, 19 17 

1 91 7. Mungos siamensis Kloss, J. N.H. Soc. Siam, 2: 215. Muang Prae, Northern 

Herpestes edwardsi GeofTroy, 181 8 Indian Grey Mongoose 

Approximate distribution of species: Arabia, Persia, Iraq, Afghanistan (Pocock); 
Baluchistan, North-\\'est Frontier, southwards over Peninsular India to Ceylon; 
Nepal and Assam. (Introduced Malay States.) 



Herpestes edwardsi edwardsi Geoffroy, 1818 

1818. Ichneumon edicardni E. Geoffroy, Descr. Egyptc, 2: 139. "East Indies" (Madras, 

Pdcock, 1933). 
1818. Ichneumon gnseus Geoffroy, loc. ctt. 157. 

1823. Herpestes frederici Desmarest, Diet. Sci. Nat. 2C): 60. Malacca. 
1829. Mangusta malaccensis Fischer, Syn. Mamm. 164. Malacca. 
l?) 184 I. Herpestes pallidus\\'as,neT, Schreb. Saugcth. Suppl. i>.- 311. 
1841. Herpestes ponliccriana Gervais, \'oy. de la Bonite, /.• 32. Pondicherry, India. 
1915. Munoos mungo ellioti Wroughton, J. Bombay X.H. Soc. 24: 52. Dharwar, 

India. Not of Blyth, 1851. 
1 92 1. Herpestes edwardsi carnaticus Thomas, J. Bombay N.H. Soc. 28: 23. Dharwar, 

For use of the name edwardsi instead of mungo see Wroughton, 1921, J. Bombay N.H. 

.Soc. 27: 547. 
Range: Peninsular India, south of the Narbada River, from Ratnagiri to Travancore 
and Madura; Eastern Ghats (Pocock). 

Herpestes ed%v.\rdsi nyula Hodgson, 1836 

1836. Mangusta (Herpestes) nyula Hodgson, J. Asiat. Soc. Bengal, j.- 236. Nepal 

iqi-,. Miingos mungo moerens Wroughton, J. Bombay N.H. Soc. 24: 52. Ganoor, 

Nimar, India. 
Range: Northern India, from Nepal to Assam, north of the Ganges; and from Cutch 
toBengal, south cil tliat river. 

Herpestes edwardsi ferrugineus Blanford, 1874 

1874. Herpestes ferrugineus Blanford, P.Z.S. 661, pi. 81. Larkhana, Sind, India. 

1884. Herpestes' andersoni Murray, Vert. Zool. of Sind, 34. Kotree, Sind. 

1 9 14. Miingos mungo palkns Ryley, J. Bombay N.H. Soc. 22: 660. Palanpur, Northern 

Gujerat, India. 
1936. Herpestes griseus montanus Bechthold, Z. Saug. //.■ 149. Hazara, Northern 

Range: Desert districts of North-Western India in valley of the Indus and Sutlcj, 
and in Rajput.uia, westwards to Baluchistan, Persia, Iraq and .\rabia. 

Herpestes edwardsi lanka Wroughton, 191 5 

1852. Herpestes griseus Kclaart, Prodr. Faun. Zeyl. 41. Not of Gcoflroy, 1818. 
1888. Herpestes munon Blanford, Mamm. Brit. India, 123, in part, not of Gmelin, 

1915. Mi/ngos lauka Wmughton, J. Bombay N.H. Soc. 24: f,;^. Cheddikulani, 177 ft., 

North Pro\incc, Geylon. 

Herpestes sxnithi Gray, 1837 Ruddy Mongoose 

Approximate distribution of species: from Rajputana and Bengal southwards to 



Herpestes smithi smithi Gray, 1837 

1837. Herpestes smithii Gray, Charlesw. Mag. N.H. /; 578. Said to be from near 

Bombay, India. 
(?) 1839. Herpestes thysanurus Wagner, Mtinch. Gel. Anz. g, 184: 439. Kashmir. 

1851. Herpestes ellioti Blyth, J. Asiat. Soc. Bengal, 20: 162. The Carnatic, India. 

1852. Herpestes torquatus Kelaart, Prodr. Faun. Zeyl. 44, nom. nud. ? Southern India. 
1864. Herpestes jerdonii Gray, P.Z.S. 550. Madras. 

1867. Herpestes jnonlicoliis jerdon, Mamm. Ind. 135. Inland from Nellore, India. 

1921. Herpestes smithii rusanus Thomas, J. Bombay N.H. Soc. 28: 25. Sambhar, 
Rajputana, India. 

192 1. Herpestes smithii canens Thomas, J. Bombay N.H. Soc. 28: 25. Mt. Abu, Raj- 
putana, India. 

Range: Rajputana, east to Bengal, southwards through Eastern and Western Ghats. 

Herpestes smithi zeylanius Thomas, 192 1 

1 92 1. Herpestes smithii zeylanius Thomas. J. Bombay N.H. Soc. 28: 24. Mankeni, East 

Province, Ceylon. 
1852. Herpestes rubiginosus Kelaart, Prodr. Faun. Zeyl. 43, not of Wagner, 1841. 

Herpestes fuscus \\'aterhouse, 1838 Indian Brown Mongoose 

Approximate distribution of species: Southern India and Ceylon. 

Herpestes fuscus fuscus Waterhouse, 1838 

1838. Herpestes fusca Waterhouse, P.Z.S. 55. India. Range: Southern India, typically 
in the hills, from 3,000 ft. to nearly 6,000 ft. (Pocock). Specimens quoted from 
Nilgiri Hills, Palni Hills, Coorg, Madura, Travancore. 

Herpestes fuscus flavidens Kelaart, 1850 

1850. Herpestes flavidens Kelaart, J. Ceylon Br. Asiat. Soc. 2: 20g (323 of 1887 reprint). 

Kandy, Ceylon. 

1851. Herpestes fulvescens Kelaart, J. As. Soc. Bengal, 20: 162. Kandy, Cevlon. 
(?) 1887. Herpestes ceylanicus Nevill, Taprobanian, /.• 62. Trincomalee, Ceylon. 
1924. Herpestes flavidens ceylonicus Thomas, Ann. Mag. N.H. i^: 240. (for ceylanicus 

1924. Herpestes flavidens phillipsi Thomas, Ann. Mag. N.H. i^: 240. Mousakande 

Estate, Gammaduwa, Central Province, Ceylon. 
Range: throughout the mountainous districts of the Central Province of Ceylon to 
over 6,000 ft., westwards to the coast near Colombo in the wet zone, and eastward 
to Uva in the dry zone; also Trincomalee. 

Herpestes fuscus maccarthiae Gray, 1851 

1851. Cynictis maccarthiae Gray, P.Z.S. 131, pi. 31. Jaffna, northern point of Ceylon. 

Herpestes fuscus sicc.\tus Thomas, 1924 

1924. Herpestes flavidens siccatus Thomas, Ann. Mag. N.H. i^: 240. Possibly .Aripo, 
near Mannar, North Province, Ceylon. 



Herpestes fuscus rubidior Pocock, 1937 

1937. Herpestes fuscus ruhidior Pocock, J. Bombay N.H. Soc. jjc).- 233. Anasigalla, 
Matugama, \\'cst Province, Ceylon. Range: South-Western Ceylon. 

Herpestes ritticollis group 

Herpestes vitticollis Bennett, 1835 Striped-necked Mongoose 

Approximate distribution of species: Western Ghats, Coorg, Travancore, etc., in 
Southern India and Ceylon. 

Herpestes vitticollis vitticollis Bennett, 1835 

1835. Herpestes vittieollis Bennett, P.Z.S. 67. Travancore, India. 

1 84 1. Crossarchus rubiginosus Wagner, Schreb. Saugeth. Suppl. 2: 329. "East Indies." 
Range: Western Ghats, Coorg, Travancore; and Ceylon. 

Herpestes vitticollis i.nornatus Pocock, 1941 

1941. Herpestes vitticollis inornatus Pocock, Fauna Brit. India, Mamm. 2: 49. Chipgeri, 
North Kanara, India. 

Herpestes urva Hodgson, 1836 Crab-eating Mongoose 

Approximate distribution of species; Fukien and Hainan, Southern China; For- 
mosa; Nepal, Assam, Burma; Indo-China, south to Peninsular Siam. 

Herpestes urva Hodgson, 1836 

1836. Gulo urva Hodgson, J. Asiat. Soc. Bengal, 5.- 238. Nepal. 

(?) 1830. Viverra fusca Gray, 111. Ind. Zool. /, pi. 5 (see Pocock, 1937, J. Bombay 
N.H. Soc. 39: 237). 

1837. Urva cancrivora Hodgson, J. Asiat. Soc. Bengal, 6: 561-4. Nepal. 

1907. Urva hanensis Matschie, W'iss. Ergebn. Exped. Filchner to China, 10, i: 190. 

Hankow, China. 
1936. Herpestes urva annamensis Bechthold, Z. Saugeth. //.• 150. Phu Qui, Annam, 

1936. Herpestes urva forrtmsanus Bechthold, loc. cit. 151. Formosa. 

1936. Herpestes urva sinensis Bechthold, loc. cit. 152. Kwantung, Southern China. 
Range: as under the species above. 

Genus ICHNEUMIA I. Geoffroy, 1837 

1835. Lasiopus I. Geoffroy, in Gervais's Resume des Le(jons do Mamm. professees au 
Mus. Paris, /.• 37. Herpestes albicaudus G. Cuvier. Not Lasiopus Dejean, 1833. 

1837. Ichneumia I. Geoffroy, Ann. Sci. Nat. Zool. 8: 251. New name to replace 
Lasiopus, preoccupied. 

I species: hhnewma alhicauda. page 298 


Ichneumia albicauda G. Cuvier, 1829 White-tailed Mongoose 

Approximate distribution of species: Southern Arabia; Ethiopian Africa, from 
Senegal to the Sudan and Somaliland, southwards to South-West Africa, the Trans- 
vaal, Natal, and Albany district. Eastern Cape Province. 

Ichneumia albicauda albicauda G. Cuvier, 1829 

1829. Herpestes albicaudus G. Cuvier, Regne Anim. ed. 2, /; 158. Senegal. 

1833. Herpestes leucurus Hemprich & Ehrenberg, Symb. Phys. Mamm. 2: h, pi. 12. 

Dongola, Sudan. 
Range: to Muscat district of Arabia. "I have no good reason for separating the 

Arabian specimens from the Sudan ones in spite of their geographical separation" 

(Morrison-Scott, 1939, Nov. Zool. 41: 198). 


Genus: Hyaena, page 299 

Genus HYAENA Brisson, 1762 

1762. Hyaena Brisson, Regn. Anim. ed. 2, 13 and 168. Canis hyaena Linnaeus. Hop- 
wood, 1947, P.Z.S. ///.• 533-536, would disregard Brisson and date Hyaena 
from Brunnich, 1771, Zool. Fundamcnta, 34, 42, 43, with type Canis hyaena 

1868. Euhyaena Falconer, Palaeontol. Memoirs, 2: 464. Canis hyaena Linnaeus. 

I species in the area covered by this list: 
Hyaena hyaena, page 299 

Hyaena hyaena Linnaeus, 1758 Striped Hyaena 

Approximate distribution of species: Transcaucasia (on west coast of Caspian Sea, 
as far north as Derbent and Dashlagar), Southern Russian Turkestan, Kopet-Dag, 
Tedshen and Atrek valleys, south of Usbekistan, south-east of Tadjikistan; Persia, 
Iraq, Syria, Palestine, Arabia; also, according to Bobrinskii, Afghanistan and Asia 
Minor; Kashmir to Nepal Terai, Baluchistan, Sind and Cutch, southwards about to 
Nilgiri Hills (perhaps further) ; Morocco, Algeria, Egypt, Libya; south of the Sahara, 
from Asben, Somaliland, Sudan and Kenya. 

Hyaena hyaena hyaena Linnaeus, 1758 

1758. Canis hyaena Linnaeus, Syst. Nat. loth ed. /.■ 40. Benna Mountains, Laristan, 

Southern Persia. 
1777. Hyaena striata Zimmermann, Spec. Zool. Geogr. 366. Renaming oi hyaena 

Linnaeus. Unavailable — Bull. Zool. Nomencl. 1950, 4: 547. 
1780. Hyena striata Zimmerman, Geogr. Gesch. 2: 256. 

1808. Hyaena orientalis Tiedemann, Zool. 350. Renaming of hraena Linnaeus. 
1820. Hyaena fasciala Thunberg, Sv. Vet. Akad. Handl. /: 59. Renaming oi hyaena 

1820. Hyena antiquorum Temminck, .\nn. Gen. Sci. Phys. Jj: 51. Renaming o( hyaena 

^' 299 

PAI.Al'.ARtniC: AND IMJIAX MAMMALS i7-,8-i946 

Hyaena hvaena hyaena [km/i/.] 

1840. Hyaena rirgata Ogilbv, in Ro\ Ic, Illustr. Hot. Himalaya, Ixvi. Rcnamine; of 

hyaena Linnaeus. 
1844. Hvaena rnlgari': indiea Blain\illc. Ostcoi;r. Manim. -•, Hyenes, 82 and cxpi. of 

pi. 6. India. 
I?) K)0^. Hvaena bokcharensii Satunin, Mitt. Kaiik. Mus. 2: 8. Bokhara, Russian 

?i 1905. Hvaena liilkieiaezi Satunin, .Mitt. Kauk. .\Iui. r.- q. .\shabad, Russian 

1905. Hvaena vuloaiii zarudnvi Satunin, Mitt. Kauk. .\Ius. 2: 14, 19. Karun River, 

South-Western Persia. 
1910. Hvaena [HvaenA] vulgaris saliinini Matsrhie, S.B. Ges. Xat. Fr. Berlin, 363. 

Range: Russian, Indian range of species, Persia, Iraq. 

Hy.-'iEN..\ hyaena vi;lgaris Desmarest, 1820 

1820. Hvaena vnlsaii^ Desmarest, Encyclop. Metli. Mainni. 215. ? Egypt. 

Hv.^E\.'\ HV.\E.\'.\ B.\RB.\R.\ Blainviilc, 1844 

1844. Hvaena rnlgari ^ harhara Blain\iile, Osteogr. Manim. Hvaena, pis. 2 and 6. Oran, 

Western Algeria. 
1853. Hvaena siiilla Filippi, Mem. R. .A.ccad. Torino, /j, 2: 131. Locality unknown. 

Hy.\e\a n\,\v.y.\ syri-\c:a Matschie, 1900 

1900. Hvaena svriaea Matschie. S.B. Ges. Nat. Fr. Berlin, 54-57. Antiochia, Syria. 

Hyae.\a hyaen,\ sultana Pocock, 1934 

1934. Hvaena hvaena sultana Pocock, .Ann. Mag. N.H. 14: 636. Mt. Q_ara, 1,500 ft., 
Ain, South-Eastern Arabia. 


Genera: Acinonvx, page 320 
Fclis, page 301 
Neqfelis, page 314 
Panthera, page 315 

Pocock split the Gats into many genera. However, the consensus of opinion is 
overwhelmingly in favour of referring most or all Ciats except Acinonvx to the Linnean 
genus Felis, or at least of recognizing as few genera as possible in this family. Simpson 
1 1 14 5, I II), 231) retains Felis, Panlhera and Acinonvx, with many subgenera of the first 
two. As this author points out, "the work of Pocock, Sonntag, Haltenorth, and others, 
shows bcvond serious doubt that the most distincti\'e group of species sometimes 
inc luded in Felis. \ensn lato, is that typified by the so-called big Gats, Lion, Tiger, 
Panther, etc., the prior name for which is Panthera. This seems to be a good genus by 
.ni\ moflern stanfl.irds". We lulK support these remarks. Pocock divided the Cats 
iiitii three sublamilies, Felinae, Pantlierinae .uid Acinonychinae, which correspond 


roughly to the three Hving genera Panthera, Felis and Acinonyx of Simpson. The genus 
or subgenus Neofelis was placed in the FeHnae by Pocock, but in the genus Panthera by 
Simpson. Judging by Pocock's figures and remarks, it is a thoroughly distinct type, 
and we are venturing to hst it as a full genus. Otherwise, we follow Simpson in 
principle, and Pocock in details of synonymy as regards various groups which he con- 
siders of generic rank (= subgeneric rank of Simpson). It may be added that the 
subgenera of Felis sensu lata are most useful in indicating the approximate position of 
a species within this large genus. 

Genus FELIS Linnaeus, 1758 

758. Felis Linnaeus, Syst. Nat. loth ed. /.• 41 . Felis catus Linnaeus, the domestic cat. 

792. Lynx Kerr, Anim. Kingd. Cat. Mamm. Nos 288-299. Felis lynx Linnaeus. Valid 
as a subgenus. 

821. Lynceus Gray, London Med. Repos. /j.- 302. Felis lynx Linnaeus. 

829. Pardina Kaup, Entw. Gesch. u. Nat. Syst. Europ. Thierwelt, /.• 53, 57. Felis 
pardina Temminck. 

834. Lynchus ]a.Td\ne, Nat. Libr. Mamm. 4: 274. Felis lynx Linnaeus. 

841. Otocolobus Brandt, Bull. Acad. Sci. St. Petersb. g: 38. Felis manul Pallas. \'alid 
as a subgenus. 

843. Chans Gray, List. Mamm. B.AL 44. Felis chaus Guldenstaedt. 

843. Caraeal Gray, List. Mamm. B.NL 46. Caracal melanotis Gray = Felis caracal 
Schreber. Valid as a subgenus. 

855. Catus Fitzinger, Wiss. pop. Nat. der Saugeth. /.• 265. Felis catus Linnaeus. 

858. Profelis Severtzov, Rev. ALtg. Zool. 10: 386. Felis celidogaster Severtzov = Felis 
aurata Temminck (the \Vest African Golden Cat). Valid as a subgenus. 

858. Cn/o/)'«.v SevertzoN-, Rcw Mag. Zool. /o: 387. /^c/« f/w!« Giildenstaedt (restrict- 
ed by Satunin, 1905). 

858. Prwnailurus Severtzov, Rev. Mag. Zool. 10: 387. Felis pardochrous Hodgson = 
Leopardus horsfieldii Gray (a race of Felis hengalensis Kerr). \'alid as a sub- 

858. ^ibethailurus Severtzov, Rev. Mag. Zool. 10: 387. Felis viverrinus Bennett. 

858. Catopiima Severtzov, Rev. Mag. Zool. 10: 387. Felis moormensis Hodgson = 
Felis temminckii \'igors & Horsfield. 

858. Pardofelis Severtzov, Rev. Mag. Zool. 10: 387. Felis marmorata XLartin. \'alid 
as a subgenus. 

858. Ictailurus Severtzov, Rev. Mag. Zool. 10: 387. Felis planiceps \'igors cS: Hors- 
field. Valid as a subgenus. 

858. Urolynchus Severtzov, Rev. Mag. Zool. 10: 389. Felis caracal Schreber. 

858. Leptailurus Se\'ertzov, Rev. Mag. Zool. 10: 389. Felis serval Schreber. \'alid as 
a subgenus. 

858. Chrysailurus Severtzov, Rev. Mag. Zool. 10: 389. Fclis nealecta Gra\- = Felis 
aurata Temminck. 

864. Serval Brchm, Fiihrer Z. Garten Hamburg, 6th ed. -^3. Serval mactilatus Brchm. 

866. Galeopardus Heuglin & Fitzinger, S.K. Akad. ^\'iss. Wien. Math. Nat. C;i. 5./, 

1 : 557. Felis serval Schreber. 

867. Viverriceps Gray, P.Z.S. 268. Felis viverrinus Bennett. 



Felis [contd.] 

1867. Cervaria Gray, P.Z.S. 276. Lvncus pardinus = Felis pardina Temminck. Not of 

\Valker, 1866. 
i86g. Aihirogale Fitzinger, S.B. Ak. Berlin, 60, i : 249. Felis planiceps Vigors & 

1870. Ailiirimis Gervais, Nouv. Arch. Mus. Paris, 6: 159. Naming of "I'Ailurin" 

Gervais, 1855, H.N. Mamm. 2: 87 (^ Felis planiceps Vigors & Horsfield). 
1874. Pvrofelis Gray, Ann. Mag. N.H. 14: 354. Felis temminckii Vigors & Horsfield. 
1885. Ailurina Trouessart, Bull. Soc. Angers, 14: Suppl. 100. Naming of "I'Ailurin" 

Gervais, 1855, H.N. Mamm. 2: 87 (= Felis planiceps Vigors & Horsfield). 
1894. Servalina Greve, Nova Acta Acad. Caes. Leop. Carol., Halle, 6^: 76. Felis serval 

1898. Oncoides Trouessart, Cat. Mamm. /.• 357. Not of Severtzov, 1858. 
1903. Eiicervaria Palmer, Science, N.S. ij: 873. Substitute for Cervaria Gray. 
1905. Tnchaelurus Satunin, Ann. Mus. Zool. St. Petersb. g: 495. Proposed as a 

substitute for Otocolobus which was thought to be preoccupied. See Pocock, 

1939, Fauna Brit. India, /.■ 315. 

1925. Poliailiirus Lonnberg, Arkiv. Zool. Stockholm, 18A, 2; 2. Felis pallida 

Buechner = Felis bieti Milne-Edwards. 

1926. Microfelis Roberts, Ann. Transvaal Mus. //.• 250. Felis nigripes Burchell, from 

South Africa. 
1926. Eremaelurus Ognev, Ann. Mus. Zool. Leningrad, 2y: 356 Eremaelurus thinobius 

Ognev (a race of Felis margarita Loche). 
1932. Badiofelis Pocock, P.Z.S. 749. Felis badia Gra), from Borneo. Valid as a 


14 species in the area covered by this list: 

Felis bengalensis, page 312 Felis margarila, page 307 

Felis bieti, page 306 Felis marmorata, page 3 1 1 

Felis caracal, page 310 Felis rubiginosa, page 314 

Felis chaus, page 306 Felis serval, page 3 1 1 

Felis libyca, page 304 Felis silveslris, page 303 

Felis lynx, page 308 Felis temmincki, page 3 1 1 

Felis manul, page 308 Felis viverrina, page 314 

Pocock, 1939, Fauna of British India, Mamm. /, keys ten of these species in some 
detail. In that work he adopted the name Constantino for the smaller species currently 
known as F. libyca, but later came to the conclusion that constantina is based on a race 
of F. serval, which he shows to occur in Algeria, and therefore he reverted to the name 
lihrca for the small African Wild Cat. In his Catalogue of the genus Felis ( 1 95 1 ) he compares 
in detail three of the Palaearctic species, silvestris, bieti and margarita (none of which 
occur in India), with their nearest alhes. For the characters of F. (Leptailurus) serval, 
see Pocock, 191 7, Ann. Mag. N.H. 20: 329-350, Classification of the existing Felidae. 

The arrangement of the species silvestris, libyca, bieti, margarita here adopted follows 
that of Pocock, Catalogue of the genus Felis. 

The nine subgenera here listed follow Pocock as far as their content of species is 
concerned. That author gave them all generic rank. In the abnve generic synonymy 
extralimital American names have not been dealt with. 



Subgenus FELIS Linnaeus, 1758 

Pocock regards the following names as synonyms oi Felis catus Linnaeus, 1758, the 
domestic cat: 

1837. Felis pulchella Gray, Mag. N.H. /; 577, Egypt, and inconspicua, loc. cit. Nepal. 
1904. Felis daemon Satunin, P.Z.S. 2: 162. Caucasus. 

1906. Felis ocreata agrius Bate, P.Z.S. igo§, 2: 317. Crete. 

Felis silvestris Schreber, 1777 European Wild Cat 

Approximate distribution of species: Scotland, Spain, France, Belgium, Italy, 
Sicily, Germany, Poland, Hungary, Yugoslavia, Rumania, Bulgaria, Greece; 
possibly still in Switzerland, Austria and Northern Portugal; Ukraine and Caucasus; 
Asia Minor. 

On this species see Pocock, 1934, J. Linn. Soc. ^ool. jg: i. 

Felis silvestris silvestris Schreber, 1777 

1777. Felis {Catus) silvestris Schreber, Saugeth. j: 397. Germany. 

1777. Felis catus ferus Erxleben, Syst. Regn. Anim. /.■ 518. 

1896. Catus ferox Martorelli, Atti Soc. Ital. Sci. Nat. Milano, jj.- 253. Lapsus for ferus. 

Range: Central Europe, from France, Northern Spain and Italy eastwards into 

South- Western Russia, western shores of the Black Sea, and probably 

Greece (Pocock). 

Felis silvestris morea Trouessart, 1904 

1904. {Felis catus) morea Trouessart, Cat. Mamm. Suppl. 273. Based on Felis catus ferus 

var. e Morea of Reichenbach, 1852, Vollstandigste Nat., Raubsaugeth. 362. 
Above Dragomanou, near Mt. Diaphorti, West-Central Morea (Pelopon- 
nesus), Greece. (Harper, 1940, J. Mamm. 21: 193.) Range: Southern 

Felis silvestris caucasica Satunin, 1905 

1905. Felis catus caucasicus SdiXMmn, Mitt. Kauk. Mus. 2: 154, 316. Borzhom, Caucasus. 
1916. Felis silvestris trapeziaBlackler, Ann. Mag. N.H. 18: 73. Khotz, near Trebizond, 

500 ft., Asia Minor. 

Felis silvestris grampia Miller, 1907 

1907. Felis grampia Miller, Ann. Mag. N.H. 20: 396. Invermoriston district, Inver- 

ness, Scotland. Range: now restricted to the wilder parts of Scotland, north 
of a line between Glasgow and Dundee (Pocock). 

Feus silvestris tartessia Miller, 1907 

1907. Felis tartessia Miller, Ann. Mag. N.H. 20: 397. Goto Doiiana, Huelva, Spain. 
Range; Southern Spain. 

Felis silvestris molisana Altobello, 1921 

1 92 1. Felis molisana Altobello, Fauna dell Abruzzo, Mamm. 55. Molise, Italy. 



Felis silvestris euxina Pocock, 1943 

i()4'V Fdis iilvestris euxina Pocock, Ann. Mag. X.H. 10: 701 . I5aspunar, in Dobrudscha, 

Felis libyca I'urster, 1780 African Wild Cat 

.\pproxiniatc distribution of species: Islands of Sardinia, Corsica and Majorca in 
Mediterranean; .South-Eastern Transcaucasia, Russian Turkestan, Kazakstan (except 
northern parts'): C^hincsc Turkestan; Arabia, Iraq, Persia, Palestine, Syria, Afghani- 
stan; Punjab, Sind, Clutch, Rajputana and Ontral India; Morocco, .\lgeria, Libya, 
Egypt; Africa,, south of the Sahara from Northern Nigeria, .\sben, the Sudan and 
Somaliland southwards to Transvaal, Natal, and regions of King Williams Tdwn 
and Cape .\gulhas in C'ape Pro\ince. 

Felis liby(:.\ lihvc^ Forster, 1780 

1780. Fdis lyhira sic) Forster in Buflbn's Nat. \'ierf. Thiere, 6: 313. Gafsa, Tuuis. 

The original spelling of this name was adopted by Pocock and G. Allen, but 

we think Forster made a mistake which comes under the heading ol a lapsus. 

It ((uild not ha\e been ignorance, since the name "I.ibya" was in cimnnon 

use by the Romans; the Latiir for Libyan is lihrcus. 
17Q2. Felix (sic) Ivnx lybiensis Kerr, Anim. Kingd. if^G. Gafsa, Tunis. 
1885. Felis crislata Lataste, .Actes Soc. Linn. Bordeaux, ^9.- 229. Not of Falconer & 

Cautley, 1836. Haidra, Tunis. 
Range: from Morocco, Algeria and Tunis to Egypt; through Nubia to the ,'\ngl(]- 
Egypti.m Sudan, and eastwards to Suakin and Massowah; and, according to 
Flower, the western coast of Sinai (Pocock). 

Felis libyc.x orn.viw Ciray, 1830 Indian Desert Cat 

1830. Felis ornata Gray, lUustr. Ind. Zool. /, pi. 2. India. 

1834. Fells serial ina yArd'me, Nat. Libr. Felinae, 4: 232. India. 

18(13. Felis tivqimta Blyth, P.Z.S. 185 (in part; not of C:u\ier, 1826, which is based on 

a feral domestic cat). 
Range: Indian range of the species, as listed abo\c. 

Felis LiHYr;.\ HtBASTis Hemprich & Ehrenberg, 1833 

1833. Felis hubastis Hemprich & Ehrenberg, Symb. Phys. Mamm. i'.- ii verso, Etjypt. 
(The Sacred Cat of ancient Egypt.) 

Felis libvi;a cai'data Gray, 1874 

1874. CluiKs eiiiiilaliis Gray, P.Z.S. 31, pi. (). Kokand, Fergana, Eastern Russian 

'Furkcstan Western AvA part of S)r-I)arya district, according lo ()giie\ ). 

? i<)i-,. FeliK eauilala ■.ehniliukovi li'nuU. .Aim. .\Ius. Zool. .-^cad. Sci. i<): 11. Kop.d 

distrii t, .Semirech\ia, Eastern Russian Turkestan. 
ii|i-|. Fells eaudata unvojhva Zukowski, Arch. .\at. Berlin, 8rj, (>: i)-,. Between west Lake 15,dkash and Ri\er C:hu, Russian Turkestan. 
! ]i)\'y Fills (iiiiilahi Inrtiiitnlis Zukowsky, hie. cit. 97. Region ol Lake B.ilk.ish, 
Riissi.m Turk(-.taii. Not of lit/inger, 18(18. 



1915. Felis caudata macrothrix Zukowsky, Arch. Nat. Berlin, 80, 10: 125. Substitute for 

longipilis Zukowsky, preoccupied. 
Range: Russian Turkestan, southwards into Persia and Afghanistan, eastwards into 

Chinese Turkestan. 

Felis libyca s.^rd.'^ Lataste, 1885 

1885. Felis libyca var. sarda Lataste, Actes .Soc. Linn. Bordeaux, ^g: 231. Sarrabus, 

1896. Felis mediterranea Martorelh, Atti Soc. Ital. Sci. Nat. Milano, jj.' 266. Sardinia. 
1906. Felis ocreata mauritana Cabrera, Bol. Real. Soc. Esp. H.N. Madrid, 6: 632. 

Mogador, Morocco. 

1920. Felis lybica cyretmrum Ghigi, Mem. R. Accad. Bologna, /.• 79. Cirene, Cyrenaica, 

(?) 1929. Felis revi Lavauden, C.R. Acad. Sci. Paris, i8g: 1023. Annes Forest, on 

border of Lake Biguglia, south of Bastia, Corsica. 
(?) 1930. Felis catus jordansi Schwarz, Zool. Anz. gi: 223. Margarita, Majorca, 

Balearic Islands. 
Range: Mediterranean islands as just listed, also Morocco, Algeria, Tunis, Libya. 

Felis libyca kozlovi Satunin, 1905 

1905. Felis {Felis) kozlovi Satunin, Ann. Mus. St. Petersb. g: 533. Oasis of Ljuktschun, 
Eastern Tianshan Mountains. 

Felis libyca murgabensis Zukowsky, 1915 

1915. Felis (Felis) murgabensis Zukowsky, Arch. Nat. Berlin, 3o, 10: 127. Tachta, on 
River Murgab, 36° N., 63° E., Afghan-Turkestan border. 

Felis libyca matschiei Zukowsky, 1915 

1915. Felis (Felis) matschiei Zukowsky, Arch. Nat. Berlin, 80, 10: 130. One hundred 

and ten versts south of Geok Tepe (38° N., 57^° E.), Transcaspia. 

Felis libyca nesterovi Birula, 191 6 

1916. Felis ornata nesterovi Birula, Ann. Mus. Zool. St. Petersb. 21, suppl. i-ii. Nachr- 

Chasasch, Lower Iraq. Ranges into Southern Persia. 

Felis libyca iraki Cheesman, 1921 

1 92 1. Felis ocreata iraki Cheesman, J. Bombay N.H. Soc. sy: 33. Kuweit, Arabia. 

Ranges into Iraq (Sheik Saad, Tigris). 

Felis libyca issikulexsis Ognev, 1930 

1930. Felis ornata issikulensis Ognev, Z. Saug. j.- 67-69. North-western shore of Lake 
Issyk Kul, Eastern Russian Turkestan. 

Felis libyca tristrami Pocock, 1944 

1944. Felis lybica tristrami Pocock, Ann. Mag. N.H. //.■ 125. Ghor Seisaban, Moab, 

1867. Felis syriaca Tristram, N.H. of the Bible, 67. .Syria. Not of Fischer, 1829. 
1895. Felis maniculata Yerbury & Thomas, P.Z.S. 547. Aden, Southern Arabia. Not 

of Cretzschmar, 1826. 



Felis bieti Milne-Edwards, 1892 Chinese Desert Cat 

Approximate distribution of species: Mongolia, Kansu, Szechuan. 

Felis bieti bieti Milne-Edwards, 1892 

1892. Felis bifti Milne-Edwards, Rev. Gen. des Sci. Pures & Appliquces, j.- 671. 

Vicinity of Tongolo and Tatsienlu, .Szechuan, China. 

1893. Felis pallida Buchner, Bull. Acad. Imp. Sci. St. Petersb. 55.- 433. Southern 

Tatung Range, Kansu, China. 
1922. Felis pallida subpallida Jacohi. Abh. u. Ber. Mus. f. Tier. u. Volkerk, Dresden, 
16, I : 9. Near Sungpan, Szechuan, China. 

Felis bieti chutucht,^ Birula, 191 7 

191 7. Felis ehiitucfila Birula, Ann. Mus. Zool. Acad. Sci. Petrograd, 21, Nouv. et 
Faits Divers, i. Nor in Province Goizso, Southern Mongolia. 

Felis bieti vellerosa Pocock, 1943 

1943. Felis bieli vellerosa Pocock, P.Z.S. 113B: 172, fig. Near Yulinfu, 4,000 ft., on 
borders of Ordos and North-Eastern Shensi, China. 

Felis chaus Giildenstaedt, 1776 Jungle Cat 

Approximate distribution of species: Eastern Transcaucasia, west coast Caspian 
Sea as far north as delta of Volga, Russian Turkestan (Southern Turkmenia, whole 
of Amu-Darya, east coast Sea of Aral, Middle and Lower Syr-Darya, Lower Chu) ; 
Chinese Turkestan, Yunnan in Western China; Asia Minor, Persia, Iraq, Syria, 
Palestine, Afghanistan; Baluchistan and Kashmir, thence southwards over Penin- 
sular India to Ceylon, eastwards to Nepal and Burma; Indo-China, Siam; Egypt. 
(? Southern Algeria, Heim de Balsac.) 

Felis chaus chaus Giildenstaedt, 1776 

I 776. Felis chaus Giildenstaedt, Nov. Com. Acad. Petrop. 20: 483. Terek River, north 

of the Caucasus. 
181 1. Felis catolvnx PaUas, Zoogr. Ross. As. /.• 23. Terek River, north of the Caucasus 

(Pocock, 1939). 
1876. Felis skawiana Blanford, J. Asiat. Soc. Bengal, ./j, 2: 49. Varkand, Chinese 

Turkestan. (For status, see Pocock, 1939, Fauna Brit. India, Mamm. /.• 

290 (footnote).) 
1898. Felis chaus typica de Winton, Ann. Mag. N.H. 2: 291. 
Range: Turkestan, Caucasus, Persia, Baluchistan, Yarkand. 

Felis chaus affinis Gray, 1830 

1830. Felis affinis Gray, Illustr. Ind. Zool. /, pi. 3. Gangootri, in Tehri Garhwal, 

Northern India. 
1836. Lynchus erythrotus Hodgson, J. Asiat. Soc. Bengal, j.- 233. Nepal. 
1844. Felis jacguemonlii Geoffrey, Jacquemont's Voy. 4: 58, Atlas, 2, pis. 2, 3. Kursali, 

8,500 ft., near Dehra Dun, Northern India. 
Range: Kashmir to Sikkim; Yunnan. 



Felis chaus kutas Pearson, 1832 

1832. Felis kutas Pearson, J. Asiat. Soc. Bengal, /; 75. Midnapore, in Bengal, about 
70 miles west of Calcutta. Range: Bengal, westwards to Cutch. 

Felis chaus nilotica de VVinton, 1898 

1898. Felis chaus nilotica de VVinton, Ann. Mag. N.H. 2: 292. Near Cairo, Egypt. 
1832. Felis riippelii Brandt, Bull. Soc. Imp. Nat. Moscou, 4: 209. Egypt. Not of 
Schinz, 1825. 

Felis chaus furax de Winton, 1898 

1898. Felis chaus furax de Winton, Ann. Mag. N.H. 2: 293. Near Jericho, Palestine. 

1902. Lyncus chrysomelanotis Nehring, S.B. Ges. Nat. Fr. Berlin, 124, 147. Jordan, 

Range includes Southern Syria, Iraq. 

Felis (?) chaus maimanah Zukowsky, 191 5 

1 91 5. Felis [Felis] maimanah Zukowsky, Arch. Nat. Berlin, 80, 10: 139. Maimana 
(36° N., 65° E.). Afghanistan. 

Felis chaus fulvidina Thomas, 1928 

1928. Felis ajjinis fulvidina Thomas, P.Z.S. 834. Kampong Tomb, Annam, Indo- 
China. Range: to Siam and Burma. 

Felis chaus prateri Pocock, 1939 

1939. Felis chaus prateri Pocock, Fauna Brit. India, Mamm. /.■ 298. Jacobabad, Sind, 
Western India. 

Felis chaus kelaarti Pocock, 1939 

1939. Felis chaus kelaarti Pocock, Fauna Brit. India, Mamm. /.• 300. Cheddikulam, 

North Province, Ceylon. Range: Ceylon and Southern India (south of the 

Kistna River). 

Felis margarita Loche, 1858 Sand Cat 

Approximate distribution of species: Southern Russian Turkestan (Kara-Kum 
Desert, region south-east of Krasnovodsk, Southern Kizil-Kum, round Termez, 
west of Bokhara); Arabia (skin in B.M. from Rub al Khali, 21° N., 55° E.), 
Sinai, Algeria, southwards to Asben. 

Felis Margarita Margarita Loche, 1858 

1858. Felis margarita Loche, Rev. Mag. Zool. 10, 2 : 49, pi. i . Near Negonga, Algeria. 
1867. Felis marginata Gray, P.Z.S. 275. 

1905. Felis ocreata marguerittei Trouessart, Caus. Sci. Soc. Zool. de France, /.• 386. 
Emendation of margarita. 

Felis margarita thinobius Ognev, 1926 

1926. Eremaelurus thinobius Ognev, Ann. Mus. Zool. Leningrad, 2y: 356, pi. 26. 
Repetek, Transcaspia, Russian Turkestan. 


l'.\LALARt;rU: AND IXUIAX MAMMALS 1758-1946 

1938. Felis margarita meinrrtzhageni Pocock, Ann. Mac;. \.H. /.■ 472. Also 1938, 
P.Z.S. 108V1: 43. EI Goica, 30' .\., Alc;crian Sahara. 

Subgenus OTOCOLOBUS Brandt, 1841. "Trichaclunn" Satunin, 1905 

Felis manul Pallas, 1776 Pallas's Cat 

Approximate distribution of species: Transcaucasia, Russian Turkestan, in part 
(.Southern Turkmenia, Lower Amu-Darya), Eastern Kazakstan, Transbaikalia. 
(E\erywhcrc rare in the U.S.S.R., according to Bobrinskii.) Zungaria (according to 
G. Allen),' Tibet, Mongolia, Western China (states of Kansu, Szechuan). Afghani- 
stan, Persia. Baluchistan, Kashmir. 

Felis manul manul Pallas, 1776 

1 77!). Felis manul Pallas, Reise. Russ. Reichs, j: 692. Jida Ri\-er, south of Lake 

Baikal, Eastern Siberia. 
1 903. Tnchaclurus manul mongolicus Satunin, Ann. Mus. Zool. Acad. Lnp. Sci. St. 

Pctersb. igo4, g.- 501. Not of Lesson, 1842. 
1907. Fdi\ manul satuni Lydekker, Game Animals hidia, 334. 
Range: northern part of range of species as gi\-en above. 

Felis m.-\nul nigripecta Hodgson, 1842 

1842. Felis tdgripectus Hodgson, J. Asiat. Soc. Bengal, //.- 276. Tibet. Range: to 

Felis manul ferruginea Ognev, 1928 

1928. (Mocolohus manul ferrugineus Ognev, C.R. Acad. Sci. U.R.S.S. 308. Mountain 
ridge of Missanev, Kopet-Dag Mountains, Transcaspia. Range: South- 
Western Turkestan, Northern Persia, Afghanistan, Baluchistan. 

Subgenus D'jVX Kerr, 1792 

Felis lynx l.inn.ieus, 17-18 European Lyn.x 

.\pproximate distribution of species: formerly in the forested parts of Europe. Still 
found in Norway, Sweden, the Baltic States, Poland and the Balkans, including 
Greece, ' .Sardinia, .Spain and Portugal. Forest zone of Russia, C'aucasus: the whole 
(il Silicria as far as and including Sakhalin, but does not occur Kamtchatka; moun- 
t.iuis of Russian Central Asia (Tarbagatai, Djungar Ala-Tau, Tianshan and 
Hissar-Alai system, Western Pamirs, Kopet-Dag). Chinese Turkestan, Tibet, Mon- 
golia, Manchuria, perhaps Chihli in China. Kashmir. Asia Minor, Persia and 
^ Palestine. Also in North America. 

Ill, IS LYNX LY.xx Linnaeus, 1758 

i7"ili. i-'iin lynx Linnaeus, .Syst. Nat. loth cd. /.• 43. Near L'psala, Sweden. 

I 7112. Lynx ruli;iiN\ Kerr, .'Vnim. Kingd. Syst. Clat. Nos. 294, 295 and p. 157 ol text. 

!7<)2. I.ynx ;'nlgari\ alha Kerr, loc . eil. Forests ofSwcflen. 



1792. Felix Lmx vulgaris melinus Kerr, Anim. Kingd. Syst. Cat. No. 2q6 and p. 157 

of text. Banks of Volga, near Kazan, Russia. 
1798. Felis borealis Thunberg, Beskrifning pa .Svenska Djur. Mamm. 14. Forests of 

Northern Sweden. 
1798. Felis kattlo Schrank, Fauna Boica, /.■ 52. Bohemia. 
1820. Felis Ivncula Nilsson, Skand. Fauna, /.• 14. Wooded and mountainous regions 

of Scandinavia. 

1824. Felis cervaria Temminck, Mon. Mamm.: 106. Asia. 

1825. Felis lupulinus Thunberg, Denkschr. k. Ak. W'iss. Munchen, 9.- 189. Northern 

1825. Felis vulpinus Thunberg, loc. cit. 192. Near Upsala, Sweden. 
1829. Felis virgala Nilsson, Ilium. Fig. Skand. Fauna, pis. 3, 4. Sweden. 
Range: European range of species excluding Iberian Peninsula, Sardinia, and the 
Caucasus; eastwards to the Yenesei, Siberia. 

Felis lynx pardina Temminck, 1824. Spanish Lynx 

1824. Felis pardina Temminck, Monogr. Mamm. /.• 116. Near Lisbon, Portugal. 

1907. Lynx pardella Miller, Ann. Mag. N.H. 20: 398. Goto Donana, Huelva, Spain. 
New name for pardina Temminck, thought to have been preoccupied bv 
pardina Oken, 1816 (unavailable). Not Felis pardella Pallas, 1784. 

Felis lynx isabellina Blyth, 1847 

1847. Felis isabellina Blyth, J. Asiat. Soc. Bengal, 16: 1178. Tibet. 

(?) 1863. Lyncus tibetanus Gray, Cat. Hodgsons Coll. B.M. 4. 

1904. Felis lynx wardi Lydekker, The Field, 104: 576. Altai Mountains. 

1904. Lynchus isabellinus kamensis Satunin, Ann. Mus. Zool. Acad. Sci. St. Petersb. g: 

13. Kam, South-Eastern Tibet. 
Range: Kashmir, Tibet, north to Tianshari and Altai Mountains, and mountains of 
Russian Central Asia, Mongolia. 

Felis lynx sardiniae Mola, 1908 

igoB. Lynx sardiniae Mola, Boll. Soc. Zool. Ital. Roma, 9.- 48. Nuoro, Sardinia. 

Felis lynx dinniki Satunin, 1915 

1915. Lynx dinniki Satunin, Mem. Cauc. Mus. Ser. A. /.• 391. Name proposed for the 
North Caucasian Lynx (see Ognev, 1935, Mamm. U.S.S.R. j.- 224). 

1905. Lynx pardina orientalis Satunin, Isvest. Kauk. Mus. 2: 166. Lenkoran, Trans- 

caucasia. Not Felis orientalis Schlegel, 1857 (a Panthera). 
1922. Lynx lynx orientalis aber. guttata Smirnov, Ann. Univ. Azerbaidjan, No. 2, 37. 

No locality. 
1922. Lynx lynx orientalis aber. virgata Smirnov, loc. cit. Not of Illiger, 181 1. 

Felis lynx wrangeli Ognev, 1928 

1928. Lynx lynx wrangeli Ognev, Rysi. Ohotnik, Nos. 5-6. (N.V.) Valley of River 

Dayeh, Hotan-Haia, Verhoiansk Mountains, Eastern Siberia. 
(?) 1922. Felix lynx var. baicalensis Dybowski, Arch. Tow. Nauk. Lwow, /.• 351, nom. 

Range: Siberia, east of the Yenesei. 


Subgenus CARACAL Gray, 1843 

Felis caracal Schreber, 1776 Caracal Lynx 

Approximate distribution of species: Russian Turkestan (only deserts of Turk- 
menia, as far north as Sea of Aral); Arabia, south to Aden, Palestine, Syria, Iraq, 
Persia, Afghanistan (according to Bobrinskii) ; Baluchistan, Punjab, Sind, Cutch, 
east to United Provinces, India; Egypt, Algeria, Morocco, and Africa south of the 
Sahara, from the Sudan, Somaliland and Asben to the Transvaal and Cape Prov ince 
(Little Namaqualand, Clanwilliam, Deelfoiitein, etc.). 

(Felis c.^RACAL caracal Schreber, 1776. Extralimital) 

1776. Felis caracal Schreber, Saugeth. pi. 1 10, text j: 413, 587, 1777. Table Moun- 
tain, Cape Town, South Africa. For discussion of type locality and author, 
see J. A. Allen, 1924, Bull. Amer. Mus. N.H. 4j: 279, and Pocock, 1939, 
Fauna Brit. India, Mamm. /.- 306. 

1843. Caracal melanoth Gray, List Spec. Mamm. B..\I. 46. Renaming oi caracal. 

Felis caracal algira Wagner, 1841 

1841. Felis caracal var. algira Wagner, Reisen in der Regenschaft Algier, j.- 76, pi. 4. 

1892. Caracal berherorum Matschie, S.B. Ges. Nat. Fr. Berlin, 114. Constantine, 

1912. Felis (Caracal) herberonim spat^i Matschie, S.B. Ges. Nat. Fr. Berlin, 61. Between 
Feriana and Tebessa, Tunis. 

1912. Caracal berherorum niedjerdae Matschie, S.B. Ges. Nat. Fr. Berlin, 62. Tunis. 

1912. Felis {Caracal) nubicus corylinus Matschie, S.B. Ges. Nat. Fr. Berlin, 63. Sup- 
posed to be from Tangier, Morocco. 

Felis caracal schmitzi Matschie, 1912 

1912. Felis [Caracal) caracal schmitzi Matschie, S.B. Ges. Nat. Fr. Berlin, 64. The Dead 

Sea region, Palestine. 
(?) 1829. Felis caracal bengalensis Fischer, Syn. Mamm. 210. Bengal. Not of Kerr, 


1912. Felis (Caracal) caracal aharonii Matschie, S.B. Ges. Nat. Fr. Berlin, 66. Mouth 
of Chabur River, on Upper Euphrates, Syria. 

(?) 1939. Caracal caracal caracal IVluUer, Pocock, Fauna Brit. India, Mamm. /.' 307. 
If the view is accepted that F. caracal dates from Schreber, 1776, with type 
locality Cape of Good Hope, then it appears that the Indian and South- 
western Asiatic race, if distinguishable, should be called schmitzi Matschie, 
which seems the first available Asiatic name. 

Range: Central India, Punjab, Sind, Baluchistan, westwards at least to Arabia and 
Palestine (Pocock). 

Felis caracal michaiclis Heptner, 1945 

194-,. Fehs (Caracal) caracal mickaelis Heptner, C.R. Acad. Sci. Moscow, ^g, 3: 230. 
Bokourdak, west of Kara Kum Desert, 60 miles north of Ashabad, Turk- 


Subgenus LEPTAILURUS Severtzov, 1858 

Felis serval Schreber, 1776 Serval 

Approximate distribution of species: Algeria, and south of the Sahara from 
Senegal, the Sudan and Somaliland, southwards to South-West Africa, Transvaal 
and Eastern Cape Province (districts near Aliwal North, East London, Grahams- 
town, etc.). 

(Felis serval serval Schreber, 1776. Extralimital) 

1776. Felis serval Schreber, Saugeth. pi. 108, text, 5.' 407, 587, 1777. Cape of Good 
Hope, South Africa. 

Felis serval constantina Forster, 1 780 

1780. Felis constantina Forster, in Buffon's Nat. d. Vierf. Thiere, 6: 313. Vicinity of 
Constantine, Algeria. For use of this name see Pocock, 1944, P.Z.S. 11^: 65. 
1829. Felis caracal algiricus Fischer, Synops. Mamm. 210. Algeria. 

Subgenus PARDOFELIS Severtzov, 1858 

Felis marmorata Martin, 1837 Marbled Cat 

Approximate distribution of species: Nepal, Sikkim, Assam, Northern Burma, 
Indo-China, Malay States, Sumatra, Borneo. 

Felis marmorata marmorata Martin, 1837 

1837. Felis marmorata Martin, P.Z.S. 1836: 108. Sumatra (see Robinson & Kloss, 

1919, J. Fed. Malay States Mus. y: 261). 
f?) 1843. Felis longicaudata Blainville, Osteogr. Mamm. Felis, 4j. 
Range: Malay States, Sumatra, Borneo; recorded by Osgood (1932) from Tonkin, 

Felis marmorata charltoni Gray, 1846 

1846. Felis charltonii Gray, Ann. Mag. N.H. 18: 211. Darjeeling, Northern India. 

1847. Felis ogilbii Hodgson, Calcutta J. N.H. 8: 44. Sikkim. 

1863. Leopardus dosul Gray, Cat. Hodgson Coll. B.M., 2nd ed., 3, nom. nud. 
1863. Felis duvaucellii Hodgson, loc. cit., nom. nud. 

Range: Indian range of species as quoted above. 

Subgenus PRO FELIS Severtzov, 1858 

Felis temmincki Vigors & Horsfield, 1827 Golden Cat 

Approximate distribution of species: Tibet, Szechuan, Yunnan to Fukien, in 

Southern China; Nepal to Assam and Burma; Indo-China, Siam, Malay States, 

PALAEARtrnt: and INDIAN MAMMALS i--,8-ir)46 

Felis temmixcki temminxki \'igors & Horsficld, 1827 

1827 Ftiis kmminckii \'igors & Horsfield, Zool. J. j; 451. Sumatra. 

i8'5i. Felis moormensis Hodgson, Gleanings in Science, j; 177. Nepal. 

1863. Felis aurata Blyth, P.Z.S. 185. Not of Temminck, 1827. 

1863. Felis niorescens Gray, Hodgson's Gat. Mamm. Nepal in B.M., cd. 2, 4. Dar- 

jccling, Northern India. 
1924. Felis tcmminckii hainsei Sowerby, Ghina J. Sci. & Arts, 2: 352. Tenguch, Snuth- 

AVestern ^"unnan, China. 

Range: Nepal to Burma, Indo-China, Yunnan, south to Malay Peninsula and 

Felis temmincki tristis Milne-Edwards, 1872 

1872. Felis Irislis Milne-Edwards, Rech. Manim. 223, pi. 31. Locality unknown. 
19114. ■^'''" semenori Satunin, .\nn. Mus. Zool. .Vcad. Sci. St. Petersb. g: 524. North- 
Eastern Szechuan, China. 

Range; Tibet, Szechuan, ? Upper Burma. 

Felis temmincki dominicanorum Sclater, 1898 

1898. Felis dominicanorum Sclater, P.Z.S. 2, pi. i. Foochow, Fukicn, China. Pocock 
and Osgood list this form as a valid race; G. Allen (1938) thought it was a 
synonym u[ tristis; the following names were also placed in the synonymy of 
/r litis: 

1908. Felis temmincki mitchelli Lydekker, P.Z.S. 433. Szechuan, China. 

1922. Felis iCatopuma) nielli Matschie, Arch. Nat. 38, A, 10: 36. Weishi, Yunnan. 
Not of Matschie, 1922 {Felis (Meofelis) mclli). 

1926. Felis temmincki badwdorsalis Howell, Prne. Biol. Soc. Washington, j^.' 143. New 
name for melli Matschie. preoccupied. 

Range: Southern China. 

Subgenus PRK ).\'AfLURUS Sc\crtzo\\ 1858 ; including ..^'Z'''''''""^"""" Scvcrtzo\-, 1858) 

Felis bengalensis Kerr, 1 792 IjCopard Cat 

.\ppro.ximate distribution of species: ,'\mur-Ussuri region ol' the Far East of 
.Siberia, Manchuria, Korea, Tsushima Island (between Korea and Japan), Formosa, 
Quelpart I., Hainan, and all the larger states of C^hina, Tibet: Baluchistan and 
Kashmir, southwards to at least Coorg and Palni Hills in Peninsular India, east- 
wards to Nepal, Assam and Burma; Indo-China, .Malay States, Sumatra, Ja\a, 
Bali. Borneo, to the Philippine Islands. 

Felis BE\r;,\LE.\sis be.ntj.xlexsis Kerr, 1 792 

1792. Felii henaalensis Kerr, Anim. Kingd. 151. Southern Bengal. 
('') 1829. Felis nipalenws Horsfield & Vigors, Zool. J. ./.• 382. ? Nepal. 
1842. Leopardiis ellioti Gray, Ann. .Mag. N.H. 10: 260. Bombay Presidency. 
18(17. Feli\ wai^ati Gray, P.Z.S. 400. Tenasserim. 


1867. Felis tenasserimensis Gray, P.Z.S. 400. Tenasserim. 

(?) 1869. Felis herschelii Gray, Cat. Cam. 28. India. 

Range: Peninsular India, Burma, Siam, Indo-China, to Yunnan, China. 

Felis bengale.nsis chinensis Gray, 1837 

1837. Felis chinensis Gray, Mag. N.H. /.• 577. Probably Canton, Kwantung, Southern 

1843. Leopardus reevesii Gray, List. Mamm. B.M. 44. China. 

1872. Felis scripta Milne-Edwards, Nouv. Arch. Mus. H.N. 7, Bull.: 92, pis. 57, 58, 

fig. I. Szechuan, China. 
1872. Felis microtis Milne-Edwards, Rech. H.N. Mamm. 221, pis. 31A, 31B, figs. 

i-ib. Near Pekin, Chihli, China. Bobrinskii lists this form as a valid race of 

euplilura, from the Siberian Far East. 
1872. Felis decolorata Milne-Edwards, Rech. H.N. Mamm. 223. Near Pekin. 
1903. Felis ricketti Bonhote, Ann. Mag. N.H. //.• 374. Foochow, Fukien, Southern 


1903. Felis ingrami Bonhote, Ann. Mag. N.H. //.• 474. Van Gin Shan Mountains, 

Northern Kweichow, China. 
1905. Felis anaslasiae Satunin, .Ann. Mus. Zool. .Acad. Imp. Sci. St. Pctersb. i[)04, g: 

528. Kam (Tibet), Kansu and North-^Vestern Szechuan, China. 
1930. Felis sinensis Shih, Bull. Dept. Biol. Sun. Yatsen. Univ. Canton, No. 4, 4. 

Chinsiu, Kwangsi, Southern China. 

Range: Chinese range of the species, apparently excepting Yunnan; Formosa. 
Recorded from Annam, Indo-China, by Osgood, who gave it specific rank. 
Poc'ock (1939, 273) appears to think that scripta (with synonyms ingrami and 
anaslasiae) is a valid race. 

Felis bengalensis horsfieldi Gray, 1842 

1842. Leopardus horsfieldii Gray, Ann. Mag. N.H. 10: 260. Bhutan, Northern India. 
1832. Felis nipalensis Hodgson, J. Asiat. Soc. Bengal, /.■ 341. Not of Vigors & Hors- 
field, 1829. 

1844. Felis pardochrous Hodgson, Calcutta J. N.H. 4: 286. Nepal. 
Range: Kashmir, Punjab, Kumaon, Nepal, Bhutan, Sikkim. 

Felis bengalensis euptilura Elliot, 1871 

1871. Felis euptilura Elliot, P.Z.S. 761. Renaming of undata Radde, preoccupied. 
1862. Felis undata Radde, Reise Ost. Sibir. 106. Not of Desmarest, 1816. Amur 
Djesa, Eastern Siberia. 

1904. Felis raddei Trouessart, Cat. Mamm. Suppl. /.■ 271. 

Felis (?) bengalensis manchurica Mori, 1922 

1922. Felis manchurica Mori, Ann. Mag. N.H. 10: 609. Near Mukden, Manchuria. 

Felis beng.\lensis trevelyani Pocock, 1939 

1939. Prionailurus bengalensis trevelyani Pocock, Fauna Brit. India, Mamm. /.■ 273. 
Near Gilgit, 5,000 ft. Range: Northern Kashmir, Upper Punjab, Southern 


palaearc;tic and Indian mammals 1758-1046 

Felis rubiginosa Geoffroy, 1831 Rusty-spotted Cat 

Approximate distribution of species: Southern India (Madras, Nellore and 
Khandala arc quoted by Pocock) and Ceylon. 

Felis rubiginosa rubiginosa Geoffroy, 1831 

1831. Felis rubiginosa I. Geoffroy, Belanger, Voy. Ind. Orient. Zool. 140. Pondi- 
cherry. Southern India. 

Felis rubiginos.a. phillipsi Pocock, 1939 

1939. Prionailurus rubiginosus phillipsi Pococic, Fauna Brit. India, Manim. /.• 278. 
Mousakanda, 3,000 ft., Gammaduwa, Central Province, Ceylon. 

Felis viverrina Bennett, 1833 Fishing Cat 

Approximate distribution of species: Ceylon, Western Ghats, Western Sind, 
Kumaon and Nepal, in India; Indo-China, Siam; Sumatra, Java. (Sclater's record 
from Formosa, quoted by Kuroda, is probably erroneous.) 

Felis viverrina Bennett, 1833 

1833. Felis viverrinus Bennett, P.Z.S. 68. India, probably the Malabar coast. 

1834. Felis himalayanus jardine, Nat. Libr. Felinae, 4: 230, pi. 24. Himalayas. 
1836. Felis viverriceps Hodgson, J. Asiat. Soc. Bengal, j.- 232. Nepal. 

1867. Viverriceps bennettii Gray, P.Z.S. 268. India. 
Range : as above. 

Felis (Ictailurus) planiceps Vigors & Horsfield, 1827, Zool. J. j.- 450. Sumatra 
(distribution: Lower Siam, Malay States, Sumatra, Borneo), has been 
recorded from Patani in Peninsular Siam, but so far as we know is extra- 
limital to the present list. 

Not certainly identifiable: Felis pardcUa Pallas, 1784, Acta Acad. Sci. Imp. Petrop. 
ij8i, 1: 281. ? Cape of Good Hope. 

Genus NEOFELIS Gray, 1B67 

1867. Neofelis Gray, P.Z.S. 265. Felis macrocelis Temminck = Felis lUardi Cuvier 
(N. nebulosa diardi, from Sumatra). 

I species: Neofelis nebulosa, page 314 

Neofelis nebulosa Griffith, 182 1 Clouded Leopard 

Approximate distribution of species: Hainan, Fukien and adjacent states in 
Southern China, Formosa; Nepal, Sikkim, parts of Burma; Indo-China, Lower Siam, 
Malay States, Sumatra, Borneo. 



Neofelis nebulosa nebulosa Griffith, 1821 

1821. Felis nebulosa Griffith, Descr. Anim. (Carn.), 37, pi. Canton, Kwantung, 

Southern China. 
1922. Felis {Neofelis) melli Matschie, Arch. Nat. 88, sect. A, 10: 35. Probably near 

Range: Southern China, Indo-China. 

Neofelis nebulosa macrosceloides Hodgson, 1853 

1853. Felis macrosceloides Hodgson, P.Z.S. 192, pi. 38. Nepal. 

1843. Felis macrocelisT'ickeW, }. Asiat. Soc. Bengal, 12: 814. Not of Temminck, 1824. 
Range: Nepal to Burma. 

Neofelis nebulosa brachyurus Swinhoe, 1862 

1862. Leopardus brachyurus Swinhoe, P.Z.S. 352, pi. 43. Formosa. Available if the 
Formosan race proves racially distinct. Kuroda, 1938, Handlist Jap. 
Mamm., calls the Formosan race diardi Desmoulins, 1823, Diet. Class, j.- 
495; probably not diardi G. Cuvier, 1823, from Sumatra. 

Genus PANTHERA Oken, 1816 

1816. Panthera Oken, Lehrb. Naturgesch. 5, 2: 1052. Felis pardus Linnaeus. 

18 16. Tigris Oken, Lehrb. Naturgesch. j, 2: 1066. Felis tigris Linnaeus. \'alid as a 

1816. Leo Oken, Lehrb. Naturgesch. 3, 2: 1070. Felis leo Linnaeus. \'alid as a sub- 

1829. -^^ Brehm, Isis (Oken), 637. Felis leo Linnaeus. 

1843. Tigris Gray, List Mamm. B.M. 40. Felis tigris Linnaeus. 

1854. Uncia Gray, Ann. Mag. N.H. 14: 394. Felis uncia Schreber. Valid as a sub- 

1868. Pardus Fitzinger, S.B.K. Akad. Wiss. Wien, ^8, 1 : 459. Felis pardus Linnaeus. 

4 species in the area covered by this list: 

Panthera leo, page 'i I g Panthera tigris, page "^18 

Panthera pardus, page 316 Panthera uncia, page 320 

Hershkowitz (1948, J. Mamm. 2g: 273, and 1949, 30.- 297) holds that all Oken's 
1 81 6 names are invalid and that his Panthera in any case would not be valid for the 
lions, tigers and leopards. J. A. Allen (1902, Bull. Amer. Mus. N.H. 16: 378) took a 
different view, and many of Oken's names, including Panthera, are in current use by 
mammalogists today. For this reason, and for general reasons explained in the Intro- 
duction, we have not discarded Panthera Oken, 18 16. But for those who do not agree 
with us, Leo Brehm, 1829, is available for the great cats. 

Pocock included lions, tigers and leopards in Panthera, but placed the ounce in a 
separate genus, Uncia. Simpson (1945) included all the above and the clouded 
leopard in Panthera. W'e take a middle view, and while following Simpson in tenta- 
tively including the ounce in the genus Panthera, we diverge from him in according 
generic rank to the clouded leopard, Neofelis. 


I'.\1.AEARC;TIC and INDIAN mammals 1758-1946 
Subgenus PANTHERA Okcn, 18 16 

Panthera pardus Linnaeus, 1758 Leopard 

Approximate distribution of species: Caucasus, Kopet-Dag Mountains (South- 
Western Turkestan) and Amur region of Eastern Siberia; Manchuria, most of the 
Larger states of China (perhaps excepting Kansu), Tibet; Asia Minor, Persia, Sinai, 
Arabia; India, from Kashmir and North-West Frontier south to Ceylon, eastwards 
to Xepal and Burma, west to Baluchistan; Indo-China, Malay States, Java, Kangean 
Islands; Morocco, Algeria, Egypt (where rare); Tropical Africa, from ? Northern 
Nigeria, Sudan and Somaliland southwards to the Cape Province, where it still 
occurs in Little Namaqualand, wilder country in the mountains near Cape Town 
(for instance, rarely to Stcllcnbosch region), Grahamstown district, etc. 

Panthera parous pardus Linnaeus, 1758 

1758. Fdis pardus Linnaeus, Syst. Nat. loth ed. /.■ 41. Egypt. 

1 8 16. Panthera vulgaris Okcn, Lehrb. Nat. 5, 2: io-)8. (Unavailable.) 

Panther.-^ p.\rdus panthera Schreber, 1777 

1777. Ft'lis panthfra Schreber, Saugeth. j: 384. Algeria. 

1832. Ft-lis palearia Cuvier, H.N. Mamm. pi. 121, text. Algeria. 

1843. Felis pardus harbarus Blainville, Osteogr. Mamm. Felis, 186, pi. 8. Algeria. 
Kfc Fisher, 1829. 

Panthera pardus fusca Meyer, 1794 

1794. Fdis fusca Meyer, Zool. Ann. /.• 394. Bengal. 

1856. Felis longicaudata Valenciennes, C.R. Acad. Sci. Paris, ^2: 1036. Not of Blain- 

\'illc, 1843. C'.eylon and Malabar coast. 
1868. Panthera aniiquorum Fitzinger, S.B. Akad. Wiss. Wien, r^S: 466. Not of Gray, 

1896. Felis pardus var. melas Pousargues, Bull. Mus. H.N. Paris, 2, 5: 181. Not of 

C'uvier, i8og. 
1904. Felis pardus chinensis Brass, Nutzbare Tierc Ostasicns, 6. 
1012. Felii pardus variegata G. Allen, Mem. Mus. Comp. Z. Harvard, 40: 235. Not of 

\\agner, 1841. Changyanghsien, Hupch (Yangtze Valley), China. Range: 

Kashmir, south to Cevlon; Burma; Szechuan to Fukicn, in Southern China. 

P.^NTHER/V pardus nimr HciTiprich & Ehrenberg, 1833 

1 833. Felis nimr Hcinprich & Ehrenberg, S>-mb. Phys. Mamm. 2: gg, pi. 1 7. (Founded 

partly on an Abyssinian skin and partK' cm .ui .Arabian one.) Arabia. Status 
not sure. 

Panthera pardus tulliana Valenciennes, 1856 

1856. Felis tulliana Valenciennes, C;.R. Acad. Sci. 42: 1039. Ninfi, 40 km. east of 
Smyrna, \Vestern Asia .Minor. Range: to Transcaucasia. 

Pa.n'thkra pardi's oriextalis Schlcgel, i8-,7 

i8-)7. Felii orientalis Schlegel, Handl. dcr Dicrkunde, /; 23. Korea. 
11)03. •''"'" '■■'/'""' Biinhote, Ann. Mag. N.H. //; 475. Amur Bay. 
Range: Knrca tn .\mur district. Eastern Siberia. 



Panthera parous pernigra Gray, 1863 

1863. Lfopardus perniger Gray, Cat. Hodgson's Coll. B.M., and ed. 3, and Preface v. 
Sikkim, 6,000-8,000 ft. Ranges to Nepal. 

Panthera p.'^rdus japonensis Gray, 1862 

1862. Leopardus japonensis Gray, P.Z.S. 262. Said to be from Japan, where the animal 
does not occur. More likely Northern China (see G. Allen, 1938, 477). 

1867. Felis fontanierii Milne-Edwards, Ann. Sci. Nat. Zool. 8: 375. Near Pckin, 
Chihli, China. 

1867. Leopardus chinensis Gray, P.Z..S. 264. (Not Felis chinensis Gray, 1837.; Moun- 
tains west of Pekin, China. Listed as a valid form (under Felis, therefore pre- 
occupied) by Bobrinskii, 1944. 

1904. Felis pardus grayi Trouessart, Cat. Mamm. Viv. Foss. 268. New name for 
chinensis Gray. 

1907. Panthera hanensis Matschie, Wiss. Ergebn. Exped. Filchner to China, /o, i: 
198. Hinganfu, China. 

1930. Panthera pardus bedfordi Pocock, J. Bombay N.H. Soc. 5^.- 323. Shangchow, 
Shensi, 3,000 ft., China. Pocock adopted the name japonensis for this race; 
see discussion in G. Allen, 1938, Mamm. China & Mongolia, /.■ 478. 

Range: Northern China, and possibly to south Ussuri region. 

Panthera pardus ciscaucasica Satunin, 19 14 

1914. Leopardus pardus ciscaucasicus Satunin, Conspectus Mamm. /.• 159. Kuban 
Province, Caucasus. 

Panthera pardus saxicolor Pocock, 1927 

1927. Panthera pardus saxicolor Pocock, Ann. Mag. N.H. 20: 213. Astcrabad, Persia. 

Range: to Baluchistan. 
1878. Felis leopardus Sclater, P.Z.S. 289. Probably Persia. Not of Schrcber, 1775. 

Panthera pardus sindica Pocock, 1930 

1930. Panthera pardus sindica Pocock, J. Bombay N.H. Soc. j^: 80. Kirthar range, 
Sind-Baluchistan border. 

Paxther.\ p.\rdus millardi Pocock, 1930 

1930. Panthera pardus millardi Pocock, J. Bombay N.H. Soc. 5./.- 316, pi. 8. Kashmir. 

Panthera parous delacouri Pocock, 1930 

1930. Panthera pardus delacouri Pocock, J. Bombay N.H. Soc. j^.- 32-,. pi. 11. Hue. 

Annam, Indo-China. 
(?) 1 9 14. Felis pardus variegata Lydekker, Rowland Wards Records, 498. Not of 

\Vagner, 1841. 

Panthera parous jarvisi Pocock, 1932 

1932. Panthera pardus jarvisi Pocock, Abstr. P.Z.S. 33; P.Z.S. 546. Sinai. 
For a review of these races, see Pocock, 1930, J. Bombay X.H. Soc. 34: 64 and 307. 


palaearc:tic: and ixdian mammals 1758-1946 

Subgenus TIGRIS Oken, 1816 (Gray, 1843) 

Panthera tigris Linnaeus, 1758 Tiger 

Approximate distribution of species: South-eastern Transcaucasia (Talysh), "it 
apparenth- does not breed there, only visiting the area from Iran" iBobrinskii), 
Southern Russian Turkestan, where rare ("in very small nuinbers on the lower Hi, 
all along the Amu-Darya ... it occasionally passes from the lower Amu-Darya to the 
lower Syr-Darya" (Bobrinskii), possibly still in small numbers on the upper Murgab 
and Atrek, in Southern Turkmenia i ; Ussuri region and middle Amur region of 
Eastern Siberia. Manchuria, Lob Xor district (Chinese Turkestan), Fukien and 
perhaps adjacent parts of Southern China, but e\'idejitly rare in other parts of 
China; Persia; most of hidia (except desert regions), and east to Assam and Burma 
(we cannot trace any reliable reference to its occurrence in Kashmir), Indo-China, 
Malay States, Sumatra, Java, Bali. 


1758. Fdis tigris Linnaeus, Syst. Nat. loth ed. /.■ 41. Bengal. 

1858. Tigris sirialus Severtzov, Rev. Mag Zool. 10: 386. Renaming oC tigris. 

1867. Tigris regalis Gray, P.Z.S. 263. Renaming oC tigris. 

Range: Kumaon and Nepal Terai, southwards to Tenasserim and Peninsulai- India, 
east to Indo-China. 


1815. Felis virgata Illiger, Abh. K. Akad. Wiss. Berlin, 98 (see also Matschie, 1897, 

S.B. Ges. Nat. Fr. Berlin, 17). Mazanderan, Northern Persia (Harper, 1940, 

J. ALamm. 21: 194). 
iq04. Felis (Tigris) tigris septeritrionalis Satunin, Priroda i Ochota, /.• 57. Twelve \'ersts 

west of Lenkoran, Talysh, Transcaucasia. 
1916. Felis tigris trabata Schwarz, Zool. Anz. ^y: 353. \'alley of Ri\-er Hi, south of 

Lake Balkash, Eastern Russian Turkestan. 
Range: Transcaucasia, through Northern Persia to Northern Afghanistan to the 
Aral Sea and Lake Balkash in Russian Turkestan (formerly to the Ob basin and 
the Altai). 

Panther.\ TIGRIS LON'GiPiLis Fitzingcr, 1868 

1868. Felii longipilis Fitzinger, S.B. Akad. \Viss. Wien. §8: 455. Amurland (type 

locality restricted by Lydekker, 1901, The great and small game of Europe, 

Western and Northern Asia, and America, 288. See also Harper, 1940, 

}. Mamm. 21: 195). 
(?) 1842. Felis mongolica Lesson, Tab). Rcgn. An. r-|0. Mongolia, iwm. mid. 
187 1. Feli\ tigris \'ar. amurensis Dode, P.Z.S. 480. 
Range: Amur and Ussuri regions of Siberia and Manchuria, possibl)' into Chihli and 


Panthera tigris coreensis Brass, 1904 

1904. Feli\ tigris cnreensis Brass, Nutzbare Tiere Ostasicns, 4. Korea. 

1915. Tigrii niikadoi Satunin, Nasa ochota, .Xn. 7, 18. 'N.V. Ognc\'s reference.) 



1925. Felis tigris mandshurica Baykov, Manchzhur. Tigr, 3. Harbin, Manchuria; and 
Felis tigris mandshurica var. mikado Baykov, loc. cit. 8. [N.V.) 

Range: Korea and Southern Manchuria, through Eastern Mongolia and Northern 
China as far as the divide between the Hwang Ho and Yangtze basins (Harper, 
1945). Possibly also the Ussuri region, as it is quoted in Bobrinskii, who does not 
give exact details. 

Panthera TIGRIS AMOYENSis Hilzheimcr, 1905 

1905. Felis tigris var. amoyensis Hilzheimer, Zool. Anz. 28: 598. Near Hankow, 

Hupeh, China. 
1929. Panthera tigris styani Pocock, J. Bombay N.H. Soc. ^5.- 531. Northern China. 

(Probably from somewhere in the latitude of the Yangtze Valley, according 

to G. Allen, 1938, 472.) 
Range: Southern China. 

Panthera Tigris LECoqi Schwarz, 19 16 

1916. Felis tigris lecoqi Schwarz, Zool. Anz. 4j: 351. Kurla district (? near Bagrash 
Kul), Lob Nor region, Chinese Turkestan. 

Subgenus LEO Oken, 181 6 (Brehm, 1829) 

Panthera leo Linnaeus, 1758 Lion 

Approximate distribution of species: Gir forest in Kathiawar, India. Formerly 
occurred in Persia and Iraq, but doubtful if any survive. Tropical Africa, from 
Somaliland, the Sudan and perhaps Senegal, south to South-West Africa and the 
Kruger National Park, Transvaal, and perhaps Zululand and Swaziland. 

Panthera leo leo Linnaeus, 1758 

1758. Felis leo Linnaeus, Syst. Nat. loth ed. /.■ 41. Constantine, Algeria. 

1826. Felis leo barbaricus Meyer, Dissert. Inaug. de Genera Felium, 6. {N.V.) 1826, 

Beytr. Anat. des Tiegers, 6. Barbary. 
1829. Felis leo barbarus Fischer, Synops. Mamm. 197. Algeria. 
1829. Leo africanus Brehm, Isis (Oken), 638. Africa. 

1867. Felis leo nigra Loche, Explor. Sci. de I'Algerie, Zool. Mamm. 35. Algeria. 
1867. Leo nobilis Gray, P.Z.S. 263. Renaming oi leo. Extinct in Algeria and Tunis 

since about 1891, and in Morocco since the ig2o's. 

Panthera leo persica Meyer, 1826 

1826. Felis leo persicus Meyer, Dissert. Inaug. de Genera Felium, 6. [N.V.) 1826, 

Beytr. Anat. des Tiegers, 6. Persia. 
1829. Felis leo bengalensis Bennett, The Tower Menagerie, i. Not of Kerr, 1792. 

Hariana, Northern India. 
1829. Leo asiaticus Brehm, Isis (Oken), 638. Asia. 

1833. Felis leo goojratensis Smec, P.Z.S. 140. Ahmadabad, Gujerat, India. 
1843. Felis leo indicus Blainville, Osteographie Mamm. Felis, atlas, pi. 6. India. 
Range: Kathiawar, India, as above. 


l'.\I.ALAKC:I IC; AM) INDIAN MAMMALS 17-,); nj-jO 

Subgenus L'A'C/A Gray, 1854 

Panthera uncia Schrcber, 1776 Ounce or Snow Lcripard 

Appruxiniati- distribution of species: Eastern Russian Turkestan, north to Altai 
Mountains ^quoted by Bobrinskii from Altai (rare), Tarbagatai (?). Dzhungar Aka- 
Tau, Tianshan system iin parts common), Alai, Zeravshan and Hissar ranges, Pamir 
(more common in Western Pamir) ). Tiijct (eastwards to Kam, according to Bobrin- 
skii; certainly as lar as Gyantse, near Lhasal and, according to Ognex', Aitvn Tag in 
Cihinese Turkestan. Kashmir. 

P.\NTHERA UN"ci.\ Schrcbcr, 1776 

177b. Fiiis uncia Schrcber, Saugeth. j.- pk 100 17761 and text, 386, 586 (1777). 

Locality unknown. 
1830. Fiiis irhi\ Ehrcnberg, .'\nn. Sci. Nat. 1^1: 3(14, 40G. Renaming iA' uncia. Altai 

i8t'). Fflis unimidis Horslield : Hodgson MS.', Aim. Mag. X.H. iG: 105. Nepal. 

Genus ACINONYX I'-mnkcs, i8j8 

1828. Acinunrx Bmokcs, Gat. Anat. Zool. Mus. J. Brookes, 16, 33. Aiinonvx vcnalor 

Brookes Ftln venatica H. Smith. 
1830. C'rnailuiiii W'agler, Nat. Svst. Amph. 30. Fclii jul<ala Si hreber. 
1841. (A-nacluius (jloger, Gemeinn. Naturgesch. /; 63. Piii (ivnailiirwi W'agler. 

I species: Acinonrx jiiba/iis, page 320 

Acinonyx jubatus Sdnebcr, 1776 Gheetah 

Ai)pn>ximatf distribution of species: Southern Turkmcnia lAtrek, Kopet-Dag, 
Tedshen and Murgab regions, rare), Persia, Arabia, Iraq and, according to Bobrin- 
skii, Afghanistan and Baluchistan. According to Bodenheimer, Transjoixlania. 
Formerly Northern India, south ol the Ganges, from Bengal to, the 
Punjal) and Sind; also C:entral India and the northern part of the Deccan; but now 
alnlo^t, il not quite, extim 1 in I'Minck!. ?I,ib\a, Egypt, where rare 
Iliiwrr, lo-o)- Morocco, Rio de ()iii. In IKipiial Africi it is less rare, and occirrs 
from ' Northern Nigeiia, the Sudan, Scmialiland, south to .South-West Alrica, the 
Kru^er National Park, 1 lansxaak and probabK' Swaziland and Zululand. 

(A( i.Ni),\v>. ]ii'..'\Tiis jLK.\ii s Schrcber, 1776. Extralimital) 

177I1. Filf, julnita Schrebii', Saugeth. ;;.■ pi. 10-, (1776), text, 392, 580 iiyjj). C..\pc 

ol Guild Hnpr. .Siiuth .\lrica. 
.■'; 1801. /'V//S i;ullald Hi inKiint, Obs. Zool. 38. .■" Egypt. Status not sure. 




1 82 1. Felis venalica Griffith, Vert. Anim. Carnivora, 93. India. 

1828. Acinonyx venator Brookes, Cat. Anat. & Zool. Mus. Joshua Brookes, 16, 33. 

(?) 1913. Acinonyx raddei Hilzheimer, S.B. Ges. Nat. Fr. Berlin, 291. Merv, Trans- 


Range: Asiatic range of the species, ? North Africa and, according to Pocock, 
probably to Somaliland. Pocock thought raddei might be valid, but it is not listed 
in Bobrinskii. 


The pinnipedes were treated as a suborder of Carnivora by Simpson (1945); 
Gregory, 19 10, The Orders of Mammals; Weber, 1928, Die Saugetiere; and Anderson, 
1947, Catalogue of Canadian Recent Mammals. Pocock regarded them as being of less 
than subordinal rank. They were regarded as a distinct order by Miller, 1923, List 
of North American Recent Mammals; Ognev, 1935, The Mammals ofU.S.S.R. and adjacent 
countries, j; G. M. Allen, 1938, Mammals of China and Mongolia, i; and Bobrinskii, 
1944, Mammals of U.S.S.R. 

The standard work on the pinnipedes as a whole is still J. A. Allen, 1880, History 
of the North American Pinnipeds, which is virtually a monograph of all species occurring 
north of the equator, and includes as well a revision of those of other seas. Keys to 
the families and genera will be found in this work, together with a detailed account 
of the nomenclatorial history of each form. A useful general work on the Otariidae 
and Phocidae is Howell, 1929, Contribution to the comparative anatomy of the 
eared and earless seals, Proc. U.S. Nat. Mus. 75, 15: 1-142. 

FAMILIES: Odobenidae, page 324 
Otariidae, page 32 1 
Phocidae, page 325 


Genera: Callorhinus, page 322 
Eumetopias, page 323 
Neophoca, page 323 
Zalophus, page 323 

J. Allen (1880) gave the following characters for the northern genera: 

Callorhinus: pelage soft, with abundant underfur; ears longer; molars 12/10; smaller 
in size; grey in colour (black when young); facial part of skull short, convex; 
molars smaller than those of Arctocephalus. 



Eumctopias: pelage harsh, laeking undcrfur; ears short; molars lo/io, the fifth pair 
separated by a long space from the fourth pair. Usually larger species; colour 
yellowish-brown (reddish-brown when young). 

^alop/ws: pelage, ears, colour, size essentially as in Eiimiiopias; molars lo/io in con- 
tinuous series. Sagittal crest very high. 

To ^alophus lie referred the Australian species Z- lobatus, which occurs in Japan 
according to Kuroda. The name cinerea Peron, 1816, antedates lobatus and is used for 
that species by Iredalc and Troughton, although J. Allen thought it was unidentifi- 
able. In recent years ^alophus has become restricted to the Californian species, and 
the name Neophoca is available for cinerea. Neophoca has a much less developed sagittal 
crest than ^alophus in British Museum material, and we consider the species cinerea 
should not be referred to ^alophus. 

Genus CALLORHINUS Gray, 1859 

1859. Callorhintis Gray, P.Z..S. 359. Fhoca ursina Linnaeus. 

1866. Arctocephalus Gill, Proc. Essex Inst. 5.- 11. Not of Cuvier, 1826. 

1892. Callotaria Palmer, Proc. Biol. Soc. Washington, y: 156. Substitute for Callor- 
hinus, assumed to be a homonym of Callirhiniis Blanchard, 1850. Kuroda 
{1938) calls this genus Otoes Fischer, 181 7, which is invalid according to 
Palmer (1904) as its type, jubata Gmelin, is composite. {Otoes Fischer, 1817, 
Mem. Soc. Imp. Nat. Moscou, 5.- 373, 445.) 

I species: Callorhimn insiniis, page 322 

Callorhinus ursinus Linnaeus, 1758 Northern Fur Seal 

Approximate distribution of species: North Pacific Ocean. Besides Western North 
America, it occurs in Eastern Siberia, Kurile Islands, Japan and Korea in winter. 
For details, sec under subspecies. 

Gallorhinus ursinus ursinus Linnaeus, 1 758 

1758. Phoca ursina Linnaeus, Syst Nat. loth ed. /.■ 37. Bering Island, off North- 
Eastern Siberia. 
1828. Otaria krachenninikoivii Lesson, Diet. Class. H.N. /j; 420. Substitute for Ursus 

marinus Stcller (1751 = Phoca ursina Linn.). Bering Sea. 
Range: "rookeries on the Gommander Islands, and a few at the southern end of 
Kamtchatka and on neighbouring islands of the Kurile group; winters on the east 
coasts of Japan" (Bobrinskii). Hokkaido, Hondo ( Kuroda). 

Callorhinus ursinus curillnsis Jordan & Clark, 1899 

1899. Callorhinus curilensis ](iTdzin & Clark, Fur Seals & Fur Seal Islands of North 

Pacific, J.- 3. Robben Island, west of Kurile Islands. 
(?) 181 I. Phoca ni^ra Pallas, Zoogr. Ross. Asiat. /.■ 107. Based apparently on a young 

Range: "rookeries (jn Seal Island (east of Sakhalin) and a few on the Kurile Islands, 
winters on the coast of Korea, reached via Peter the Great Bay" (Bobrinskii). 



Genus EUMETOPIAS Gill, 1866 

1866. Eumetopias Gill, Proc. Essex Inst. 5; 7. Arctocephalus monteriensis Gray = Phoca 
jubata Schreber. 

I species: Eumetopias jubata, page 323 

For a discussion of the nomenclature of this species, see J. A. Allen, 1902, The 
names of some of the Otariidae, Bull. Amer. Mus. N.H. 16: 1 1 1. 

Eumetopias jubata Schreber, 1776 Steller's, or Northern Sea-lion 

Approximate distribution of species: North Pacific Ocean. Besides Western North 
America, occurs off Eastern Siberia ("the best-known rookeries are in the Sea of 
Japan, near Vladivostock, in the Sea of Okhotsk on loniu Island and the Yamskie 
Islands, and in Bering Sea on Cape Shipunskii (South- Western Kamtchatka)" 
(Bobrinskii) ; and Japan (recorded from Sakhalin, Kuriles, Hokkaido, N. Hondo and 

Eumetopias jubata Schreber, 1776 

1776. Phoca jubata Schreber, Saugeth. 3: 300, pi. 83B. North Pacific Ocean (eastern 

coast Kamtchatka, according to Ognev). 
1811. Phoca leonina Pallas, Zoogr. Rosso-Asiat. /.• 104. Not of Linnaeus, 1758. 
1828. Otaria stellerii Lesson, Diet. Class. H.N. i^: 420. 

Genus ZALOPHUS Gill, 1866 

1866. Zalophus Gill, Proc. Essex Inst. 5.- 7, 11. Otana gillespii MacBain = Otaria 
calif orniana Lesson. 

I species: ^alophus californianus, page 323 

Zalophus californianus Lesson, 1828 Californian Sea-lion 

Approximate distribution of species: Western North America. Recorded from the 
Kurile Islands by Kuroda ( 1 938) under the name Eumetopias gillespii. As gillespii is the 
type species of Zalophus, this author, who retains the genus ^alophus in his list for 
another species, could not have been correct in listing this form under Eumetopias. A 
specimen in the British Museum is labelled Japan. 

Zalophus californianus Lesson, 1828 

1828. Otaria californiana Lesson, Diet. Class. H.N. 13: 420. California. 

1858. Otaria gillespii MacBain, Proc. Edinb. Roy. Phys. Soc. /.- 42 2. California. 

(?) 1866. Otaria japonica Peters, Mber. Preuss. Akad. Wiss. 668. Japan. 

1874. Eumetopias elongatus Gray, Proc. Zool. Soc. Lond. 1873: 766. Japan. 

Range: Southern Mexico to Northern California, casually to British Columbia 
(Anderson). ? Japanese seas. 


l'.\l,Al.ARt:rK: AM) IMJlAN MAMMALS 1 7 j(i i.j4t, 


Genus : Odohcniis, page 324 

Genus ODOBENUS Brisson, 1762 

1762. Udobenus Brisson, Rc£;n. Anim. ed. 2, 30. Odobenus Brisson = Phoca rosmarus 

1768. Trichechus Linnaeus, Syst. Nat. r2th cd. /.• .jq. Not of Linnaeus, 1758, which is 

the Manatee. 
1772. Rosmarus Brunnich, Zool. Fundamenta, 34, 38-39. Phoca rosmarus Linnaeus. 

Hopwood, K)47, P-Z-'S- 533-536, would disregard Brisson and call this genus 
Rosmarus Brininich. However, Odobenus was adopted by Miller, Ognev, Simpson and 
virtually all recent authors, who use Brisson's names. It is hoped that the Inter- 
national Clommission on Zoological Nomenclature will endorse generic names 
dating from Brisson, 1762, since considerable confusion will be caused if they are all 

I species; Odobenus rosmarus, page 324 

Odobenus rosmarus Linnaeus, 1758 Walrus 

Approximate distribution of species: .\rctic regions of Eurasia and North ,\merica. 
Has been recorded from the Orkneys, Hebrides and Scotland (where rare) ; Holland, 
Denmark, Norway, Sweden. According to Bobrinskii it survives in small numbers in 
the Spitzbergen .\rchipelago and the Franz Joseph Islands, rarely off Iceland, coasts 
of Barents Sea, off Novaya Zemlya, in Kara Sea, Laptev Sea, Severnaya Zemlya, 
Ghukotskoe Sea and extreme north of Bering Sea, as far cast as Kamtchatka Penin- 
sula. It is cjuotcd from Japan by Kumda (Hokkaido and recorded Hondo). 

Odobe.nls rosmarus rosmarus Linnaeus, 1758 

17-yS. Phoca rosmarus Linnaeus, Syst. Nat. loth ed. /: 38. North Atlantic (Thomas, 
ii|iii. Range: from C'anada and Greenland eastwards to Novosibirskie 
^ loi I. Rinmani\ urctnin I'allas, Zoogr. Ross. ,\siat. /.• 269. Novaya Zemha. Status 
full' ()gnc\-. 



Odobenus rosmarus divergens Illiger, 1815 

1815. Trichechus divergens Illiger, Abh. Akad. \Viss. Berlin, 180^-11: 68. About 35 

miles south of Icy Cape, Alaska. 
(?) 1815. Trichechus obesus Illiger, loc. cit. 64, nom. nud. 
1 83 1. Trichechus cookii Frcmery, Bijdrag. Nat. Vetensk. 6: 385. Off Icy Cape, Alaska 

{70°o6'N., i63°i8'W.). 
(?) 1922. Trichechus orientalis Dybowski, Arch. Tow. Nauk. Lwow, /.• 351, nom. nud. 
Range: Alaska to Eastern Siberia (Bering Sea). Bobrinskii calls the eastern race 0. r. 

arcticus Pallas, 1 8 1 1 , which antedates divergens, but is a synonym of the typical race 

according to Ognev. 

For North American range of this and the typical race, see .\nderson, 1947, Cat. 
Canadian Rec. Mamm. 81. 


Genera: Cystophora, page 333 
Erignathus, page 331 
Halichoenis, page 332 
Monachus, page 332 
Phoca, page 327 

The subfamilies here admitted follow Simpson. 

All species of this family here dealt with occur in the U.S.S.R., and we include a 
translation (slightly modified) of a key to these species which Bobrinskii gives. 

Key to the species of Phocidae, from Bobrinskii, 1944, Mammals ofU.S.S.R. (Even 
if Russian is not understood, it is helpful to use this translation in conjunction with 
the original, since Bobrinskii gives text figures illustrating the characters referred to.) 

1. There is a skin pouch on the upper side of the snout which can be inflated, attain- 

ing a large size in males. Only one incisor each side in bottom jaw. Th'^ 
premaxillae clearly not reaching the nasals. CYSTOPHORA CRISTATA 
No such pouch on upper side of snout. Two incisors each side of bottom jaw. 
Premaxillae reach the nasals. 2 

2. End of nose covered with hair right up to the nostrils. Claws comparatively small, 

and on hind limbs frequently missing. Two incisors each side in top jaw. 

End of nose hairless or almost so. Claws well developed, and in the forelimbs large 
and powerful. Three incisors each side in top jaw. 3 

3. Snout very long, so that the distance between tip of nose and eye is almost twice 

that between the eye and the auditory meatus. Profile of parietals, frontals and 
nasals form a straight line. HALICHOERUS GRIPUS 
Snout much shorter, so that distance between end of nose and the eye is a good 
deal less than twice that between the eye and auditory meatus. Profile of 
parietals, frontals and nasals convex. a 



4. In the forclimbs the longest finger is the third. Vibrissae dense and straight. 

Length of aduhs, over 2 m. Four mammae in the female. Jugal short and deep, 
the depth of the bone not less than half its length. 

The third finger of fore flippers is shorter than the first or second. Vibrissae sparse, 
and wavy. Body not more than 2.2 m. long. Two mammae in the female. Jugal 
long and narrow, so that depth of the bmie is less than half its length. (Genus 

5. The bony nasal septum reaches, or almost reaches, the rear edge of the bony 

palate. Rear edge of bony palate forms a more or less straight line or shallow 
double arch. Adults, with the exception of some females that have just reached 

sexual maturity, not spotted, but with large dark and light areas. 6 

The bony nasal septum falls far short of rear edge of bony palate. Rear edge of 
bony palate forms a high arch, usually pointed at the top. Clolouring: generally 
there are small spots, less frequently the uniformly dark back gradually lightens 
towards the abdomen. 7 

6. The forclimbs and neighbouring parts of body dark, never any small dark spots on 

body. Condylobasal length of skull under 200 mm. Bony nasal septum just fails 
to reach rear edge of bony palate. The upper toothrow is curved, seen from 
below and from the side (in other words, curved in the horizontal and vertical 
planes). PHOCA F ASCI AT A 

The forclimbs and neighbouring parts of body are light-coloured, and the body 
sometimes covered with small dark spots; condylobasal length of skull in adults, 
over 200 mm. The bony nasal septum reaches the rear edge of the bony palate. 
Upper toothrow not curved. PHOCA GROENLANDICA 

7. The nasal bones are short and broad, the teeth large and the infraorbital foramen 

small, its diameter two-thirds to one-third that of the alveolus of the canine 
tooth (N. .Siiiirnos). Dominant type of colouring: small dark spots on a light 
background. PHOCA VITULINA 
The nasal bones are narrov/ and longer, the teeth small and the infraorbital 
foramen well developed, of approximately the same diameter as the alveolus of 
the canine tooth or even larger (N. Smirnov). C;olouring different (ring-spots, 
large spots or uniform colour). 8 

8. Zygomatic arches set in such a way that they cannot be seen when the skull is 

looked at from behind, being hidden by the brainpan. Adults usually spotted. 

Zygomatic arches wide .set, so that thev are easily visible when the skull is looked 
at from the back. C^ohiur usualK- unilunu, lightening towards the abdomen, 
without spots. ' ' PHOCA SIBIRICA 

t). Infraorbital foramen the same size as aKcnlus of canine tooth. Anterior nasal 
opening ( (iniparati\ely wide. Adults ring-spotted. PHOCA HISPIDA 

Infraorbital foramen wider than alveolus of canine tooth. Anterior nasal opening 
relatively narrow. Colouring: usually dark, comparatively large spots on a light 
ba.k,L;r..und. PHOCA CASPICA 



In addition, it may be added that, according to Ognev and as figured by Bobrinskii, 
the interorbital width is very narrow indeed in the subgenus Pusa {P. hispida and 
allies) and much less so in the subgenus Phoca [vitulina). 

Subfamily P h o c i 

Genus PHOCA Linnaeus, 1758 

1758. Phoca Linnaeus, Syst. Nat. loth ed. /; 37. Phoca vituUna Linnaeus. 

1777. Pusa Scopoli, Introd. Hist. Nat. 490. Phoca foetida Fabricius = Phoca hispida 

Schreber. Valid as a subgenus. 
1826. Callocephaliis F. Cuvier, Diet. Sci. Nat. ^g: 544. Phoca vitulina Linnaeus. 
1844. Pagophilus Gray, Zoology o^ Erebus and Terror, 3. Phoca groenlandica Erxleben. 

Valid as a subgenus. 
1864. Halicyon Gray, P.Z.S. 28. Halicyon richardii Gray = Phoca vitulina richardi, from 

1864. Pagomys Gray, P.Z.S. 31. Phoca foetida = Phoca hispida Schreber. 
1866. Haliphilus Gray, Ann. Mag. N.H. ly: 446. Halichoerus antarcticus Peale 

= Phoca pealei Gill ? = Phoca vitulina richardii Gray. 
1873. Histriophoca Gill, Amer. Nat. y: 179. Phoca fasciata Zimmermann. Valid as a 

1904. Pagophoca Trouessart, Cat. Mamm. Suppl. 287. Substitute for Pagophilus Gray. 

Our listing of this genus follows Miller, Simpson and others. It may be noted, 
however, that Ognev listed Histriophoca and Pagophoca as full genera. These two seals 
are strongly differentiated from the more typical subgenus and Pusa. Bobrinskii 
(1944) introduces a new arrangement, in which Pusa is synonymous with Phoca sensu 
stricto, and Pagophoca is synonymous with Histriophoca, the latter being considered as a 
subgenus of Phoca, so that according to that author's views there are two subgenera 
only in Phoca, each with two species ( or specific groups, as he keeps the Baikal and 
Caspian Seals specifically distinct from P. hispida). 

Our own view is that on account of the difference in palatal structure between the 
two main divisions in the genus, it might be possible to follow Bobrinskii's arrange- 
ment provided Histriophoca (Pagophoca included in it) were given generic rank. How- 
ever, we here adopt the customary arrangement. Bobrinskii states that P. caspica and 
P. sibirica are very close to P. hispida, possibly merely subspecies of it. 

6 species in the Palaearctic: 

Phoca caspica, page 330 
Phoca fasciata, page 330 
Phoca groenlandica, page 330 
Phoca hispida, page 328 
Phoca sibirica, page 330 
Phoca vitulina, page 328 


PAI.AKARCTK: and IXUIAX mammals 1738-1946 
.Subj;cnus I'/KX.'A Linn, tens, 1758 

Phoca vituHna Linnaeus, 1758 Common Seal 

Appioximali- dislribution of species: Sandy roasts from Spain and British Isles 
includins Ireland alont;- North-Western European coasts (France, Germany, Hollancf) 
III Denmark, Xorvvay, the Baltic, Russia (including; Novaya Zemlya, Murman coast 
where rare), Barents Sea). Eastern Siberia 1 Chukotskoe Sea, Berinc; Sea, Okhotsk 
Sea). Japan, Korea. \'arious parts of North America, Greenland included 1 for 
details see .Anderson, 1947. Canadian Recent Mammals', 78). 

Phoc.\ vitulina vitl;li.\a Linnaeus, 1758 

1758. Phoca vitulina Linnaeus, Syst. Nat. loth ed. /.• ;^8. Gulf of Bothnia, Xurthnn 

Baltic (Thomas, 191 1). ^ Where the animal does not now occur ! Bobrinskii).) 
181 I. Phoca canina Pallas, Zoogr. Ross. Asiat. /.• i i.],. 
1820. Phoca variei^ula Nilsson, Skand. Faun. /.■ 3-,9. New name for Phoca viliilitia 

1824. Phoca H'o/nilicola I'hienemann, Nat. Bcmcrk. Rcisc Europa, /.• 39, pi. 5. 

1824. Phoca littorca Thienemann, loc. cit. ? Northern Russia. 
1828. Phoca linnaci Lesson, Diet. Class. H.X. /j.- 415. 
1828. Phoca thienemannii Lesson, Diet. Class. H.N. i^: 414. New name for P. scopuli- 

cola Thienemann. 
Range: European range of species. 

Phoca vrruLL\.\ largh.^l Pallas, 181 1 

181 I. Phoca largha Pallas, Zoogr. Ross. Asiat. /; i 13. Eastern part of Kamtchatka. 

1828. Phoca chorisii Lesson, Diet. Class. H.N. ij: 417. Kamtchatka. 

1844. Phoca nummularis Temminck, Fauna Japon. 3. Japan. 

(?) 1864. Ilalicvon richardii Gray, P.Z.S. 28. Vancouver Island, British Columbia. A 

svnonym, according to Ognev. Queried by G. Allen as occurring on eastern 

Cihinese coasts. 
1902. Phoca ocholensis ]. Allen, Bull. Anirr. Mus. .\.H. i/i: 480. Not rif Palias. 181 1. 

Mouth iif Gichiga Ri\er, Okhcitsk Sea, Eastern Siberia. 
1902. Phoca ochotenui macroden^ ]. Allen, 15ull. .\mer. Mus. N.H. 16: 483. ,\\atcha 

Bay, Kamtchatka. 
1902. Phoca Ucjiicaeii ]. Allen. Bull. .Xmcr. Mus. X.H. rlj: 48-,. Bering Island, Eastern 

1935. Phoca vilnUna largha natio /lalluMi .Xauninv lK: Smirno\', Inst. Fish. 

Oce.uiogr. .Moscow, jj.' 177. Sea (if (Okhotsk. 
1041. Phoca /leleni Mohr, Zool. Anz. Leipzig, /7';.' 41). C'nast ot Korea. 
Range: Eastern Siberia, Japan, Korea, apparent) \ Western Xnrth .\merica. 

Subgenus PUSA Sciipoli, 1777 

Phoca hispida Si hreber, 1775 Ringed Seal 

,\ppr(ixim.ite distribution of species: .Xnrlhirn Europe, U.S.S.R. eastwards to 

Sakhalin anrl J.ipan, and .Arctic North .America (for some details see Anderson, 1947, 



Canadian Recent Mammals, 79) . Russian localities include the White Sea, Bering Sea, 
Sea of Okhotsk and Tatarsk Strait (also, according to Ognev, Taimyr Peninsula and 
New Siberian Islands); also the Baltic Sea, including Gulfs of Bothnia and Finland 
(it swims up the Neva to Leningrad), Lake Ladoga and some Finnish lakes (Lake 
Saima and others near it). Has been recorded from Novaya Zemlya, Iceland, Spitz- 
bergen; rare visitor to France, Germany, Denmark, Holland, British Isles (recorded 
from Norfolk and several places in Scotland) ; Norway. 

Phoca hispida hispida Schreber, 1775 

1775. Phoca hispida Schreber, Saugeth. jj.- pi. 86 (te.xt, 1776, j.- 312). Coasts of Green- 

land and Labrador. 

1776. Phoca foetida Fabricius, MuUer, Zool. Danicae Prodr., viii; 1780, Fauna 

Groenlandica, 13. Greenland. 

1820. Phoca annellata Nilsson, Skand. Faun. /.■ 365. New name {or foetida Fabricius, 

(?) 1921. Piisa hispida pygmaea Zukowsky, Arch. Naturgesch. 8yA, 10: 183. ? Green- 
land and Novaya Zemlya. 

Phoca hispida botnica Gmelin, 1788 

1788. Phoca vitulina botnica Gmelin, Linn. Syst. Nat. 13th ed. /.• 63. Gulf of Bothnia, 

Baltic Sea. 
1839. Phoca communis var. octonata Kutorga, Bull. Soc. Nat. Moscow, 185, 189. No 

1839. Phoca communis var. undulata Kutorga, Bull. Soc. Nat. Moscow, 185, 191. No 


Phoca hispida ochotensis Pallas, 181 1 

181 1. Phoca ochotensis Pallas, Zoogr. Ross. Asiat. /.• 117. Northern part of Okhotsk 
Sea, between Tamis Bay and Gichiga, Eastern Siberia. 

1902. Phoca (Pusa) hispida gichigensis J. Allen, Bull. Amer. Mus. N.H. 16: 478. 
Gichiga, Okhotsk Sea, Eastern Siberia. 

Phoca hispida saimensis Nordquist, 1899 

1899. Phoca foetida var saimensis Nordquist, Acta Soc. Fauna Flor. Fenn. 75, 7: 28. 
Lake Saima, Finland. 

Phoca hispida ladogensis Nordquist, 1899 

1899. Phoca foetida var. ladogensis Nordquist, Acta Soc. Fauna Flor. Fenn. /j, 7: 
Lake Ladoga (Finnish-Russian border). 


Phoca hispida pomororum Smirnov, 1929 

1929. Phoca hispida pomororum Smirnov, C.R. Acad. Leningrad, 95. Barents Sea; west 

coast Novaya Zemlya. 
1929. Phoca hispida pomororum natio rochmistrovi Smirnov, loc. cit. 95. Sumski Posad, 

western coast of White Sea, Northern Russia. 



Phoca hispida birulai Smirnov, 1929 

1929. Phoca hisfiida hirulai Smirnov, C.R. Acad. Leningrad, 96. New Siberian Islands; 
Liakhov Island. 

Phoca hispida krasche.ninikovi Naumov & Smirnov, 1935 

1935. Plwca hispida kraicheninikori Naumov & Smirnov, Trans. Inst. Fish. Oceanogr. 
Moscow, 5.- 182. Bering Sea, Eastern Siberia. 

Phoca caspica Gmclin, 1788 Caspian Seal 

.\pproximate distriburion of species: Caspian Sea, "distributed all over the Caspian 
Sea but collects in dilFerent parts of it according; to the time of year" ( Bobrinskii). 

Phoca caspic,\ Gmelin, 1788 

1788. Phoca vitulina var. caspica Gmelin, Syst. Nat. 13th ed. /.■ 64. Caspian Sea. 

Phoca sibirica Gmelin, 1788 Baikal Seal 

Approximate distribution of species: Lake Baikal, Eastern Siberia. 

Phoca sibirica Gmelin, 1 788 

1788. Phoca vitulina var. sibirica Gmelin, Syst. Nat. 13th ed. /.■ 64. Lakes Baikal and 

1873. Phoca baicaleiisis Dybowski, Arch. Anat. Physiol. Lpz. 109. Lake Baikal. 
1922. Phoca oronensis Dybowski, Arch. Tow. Nauk. Lwow, /.• 352, nom. nud. Lake 

Oron (right bank of Witim, Govt, of Yakutsk, about 57|° N., ii7°E.). 

(According to Ognev (1935) there is no seal in this lake.) 

Subgenus HISTRIOPPIOCA Gill, 1873 

Phoca fasciata Zimmermann, 1783 Ribbon Seal 

Approximate distribution of species: Kurile Islands, Tatarsk Strait, Sea of 
Okhotsk, Bering Sea and Chukotskoe Sea, penetrates into eastern part of East 
Siberian Sea; everywhere rare (Bobrinskii). To Alaska. Has been recorded from 
Hokkaido (Kuroda). 

Phoca fasciata Zimmermann, 1783. 

1783. Phoca fasciata Zimmermann, Geogr. Gcsch. 5.- 277. Kurile Islands, north of 

1 83 1. Phoca equcstris Pallas, Zoogr. Ross. As. /.• iii. 

Subgenus PAGOPHILUS Gray, 1844 

Phoca groenlandica Erxleben, 1777 Harp Seal (Greenland Seal) 

.\pproximatr distribution of species: Northern Europe, Russia, Western Siberia 

and northern .Xnrth America fsee Anderson, 1947, Canadian Recent Mammals, 79, for 



Nearctic range). Iceland, Spitzbergen, Jan Meyen Island districts, White Sea, Kara 
Sea, Cheshskaya Bay (Northern Russia) ; rare wanderer to British Isles, France and 
Holland. The Eastern Siberian limit is Severnaya Zemlya (Bobrinskii) . Range 
includes Norway. 

Phoca groenlandica groenlandica Erxleben, 1777 

1777. Phoca groenlandica Erxleben, Regn Anim. /.• 588. Greenland and Newfound- 


1785. Phoca semilunaris Boddaert, Elench. Anim. 170. Greenland, Iceland. 

(?) 1822. Phoca albicauda Desmarest, Mamm. 541. No locality. 

(?) 1824. Phoca leucopla Thienemann, Nat. Bemerk. Reise Europe, /; 102, pi. 13. A 
few miles north of Grimsey Island, north of Iceland. Thienemann says that 
the type specimen of kucopla was found in a herd of several hundred Phoca 
groenlandica and thinks it was just an individual variation. 

1 85 1. Phoca albini Alessandrini, Mem. R. R. Accad. Bologna, 2: 158. 

Phoca groenlandica oceanica Lepechin, 1778 

1778. Phoca oceanica Lepechin, Acta Ac. Petrop. 1777, i: 259, pis. 6 and 7. White 

Sea, Northern Russia. 
181 1. Phoca dorsata Pallas, Zoogr. Ross. As. /.■ 112. 

Genus ERIGNATHUS Gill, 1866 
1866. Erignathus Gill, Proc. Essex Inst. 5.- 5, 9. Phoca barbata Erxleben. 
I species: Erignathus barbatus, page 331 

Erignathus barbatus Erxleben, 1777 Bearded Seal 

Approximate distribution of species: Northern Eurasia, east to Sakhalin and 
rarely Hokkaido, Japan. North America, from Bering Sea to Greenland. Said to have 
been recorded from Norfolk, England, and from Scotland (River Beauly) ; Norway. 
In U.S.S.R., White Sea, all along the European and Asiatic coast of the Arctic 
Ocean, off all the islands of the Arctic Ocean, and in the Bering Sea and Sea of 
Okhotsk as far south as Tatarsk Strait; it sometimes swims a few kilometres up rivers 
(Bobrinskii). Iceland, Spitzbergen, Franz Josef Land, Jan Mayen Island. 

Erignathus barbatus barbatus Erxleben, 1777 

1777. Phoca barbata Erxleben, Syst. Regn. Anim. /; 590. Type locality restricted to 

Southern Greenland by Ognev, 1935. 

1778. Phoca leporina Lepechin, Acta Ac. Petrop. lyyj, i: 264, pi. 8. \\'hite Sea. 
1828. Phoca parsonsii Lesson, Diet. Class. H.N. 13: 414. Northern Seas. 

1828. Phoca lepechenii Lesson, loc. cit. 415. Renaming of leporina. 

Erignathus barb.\tus nauticus Pallas, 1 8 1 1 

181 1. Phoca nautica Pallas, Zoogr. Ross. As. /.■ 108. Okhotsk Sea, Eastern Siberia. 
181 1. Phoca albigena Pallas, loc. cit. 109. Kamtchatka. 



Genus HALICHOERUS Xilsson, 1820 

1820. Halichocrus Xilsson, Skand. Fauna, Dagg. Djur. /; 376. Halkhoerus griseus 
Xilsson = Phoca grvpiis Fabricius. 

I species: Halichocrui grxpiis, page 332 

Halichoerus grypus Fabricius, 1791 Grey Seal 

Approximate distribution of species: Europe, from British Lsles northward, Russia 
and in Xorth America (for American range see Anderson, 1947, Canadian Recent 
Mammals, 80). Novnya Zemlya, Barents Sea, Murman coast, neck of White Sea, 
Baltic Sea (including Finland, Gulf of Bothnia), Norway, England (rocky parts of 
west coast), Scotland, Ireland, Orkneys, Shetlands, Hebrides, Faroe Islands, Scilly 

Halichoerus grypus Fabricius, 1791 

1791. Phoca grypus Fabricius, Skrlvter af Xaturhist. Selskabet, Copenhagen, /, 2: 

167, pi. 13, fig. 4. Greenland. 
1820. Halichoerus griseus Xilsson, Skand. Fauna, Dagg. Djur. /.■ 377. Greenland. 
1824. Phoca halichoerus Tliicncmann, Xat. Bemrrk. nordl. Reise Europa, /: 142. 

1 85 1. Halichoerus macrorhynchus Hornschuch & Schilling, Arch. Naturgesch. ly, 2: 

28. Baltic Sea. 
1 85 1. Halichoerus pachvrhvnchus Hornschuch & Schilling, loc. cit. Baltic Sea. 
1886. Halichoerus grypus var. atlantica Xehring, S.B. Ges. Nat. Fr. Berlin, 122. \Vest 

coast of Norway. 
1886. Halichoerus grypus var. tallica Nehring, loc. cit. Baltic. 

SuBF.»iMiLY M o n a c h i n a e 

Genus MONACHUS Fleming, 1822 

1822. Monachus Fleming, Philos. Zool. 2: 187 (footnote). Phoca monachus Hermann. 

1824. Pelagios F. Cuvicr, Mem. Mus. H.N. Paris, //.■ 196. Phoca monachus Her- 

1841. Pelagocvon Gloger, Gemeinn. Naturgesch. /, xx.\iv, 163. Pelagocyon monachus = 
Phoca monachus Hermann. 

1848. Rigoon Gistl, Xat. Thierr. lur hohcrc Schulen, x. Xew name for Pelagios F. 

1854. Heliophoca Gray, Ann. Mag. X.H. 13: 201. Heliophoca atlantica Gray = Phoca 
monachus Hermann. 

I species in the area covered by this list: 
Monachus monachus, page 333 



Monachus monachus Hermann, 1779 Monk Seal 

Approximate distribution of species: Atlantic (Madeira, Canaries and Southern 
Rio de Ore); Mediterranean, formerly most coasts but now restricted to parts of 
Morocco, Cyrenaica, Corsica, islands in the Southern Adriatic and off Greece, Crete, 
? Egypt, Palestine and the Lebanon; Black Sea fCape Kaliakra in Rumania and 
Sosopolis in Bulgaria, and perhaps the eastern shore). 

Monachus monachus Hermann, 1779 

1779. Phoca monachus Hermann, Beschaf. Berlin Ges. Naturf Freunde, ^; 501, pis. 

12, 13. Mediterranean Sea. 
1785. Phoca alhiventer Boddaert, Elench. Anim. 170. Adriatic Sea. 
1800. Phoca bicolor Shaw, Gen. Zool. /, 2: 254. Adriatic Sea. 
1816. Phoca leucogaster Peron & Lesueur, Voy. aux Terres Austr. 2: 47 (footnote). 

Nimes, Southern France. 
1828. Phoca hermannii Lesson, Diet. Class. H.N. i^: 416. Adriatic Sea. 
1838. Monachus mediterraneiis Xilsson, K. Svenska Vet. Ak. Handl. 183J: 238. 

Adriatic Sea and Greek Archipelago. 
(?) 1843. Phoca isidorei Lesson, Echo ^Iondc Sa\ant, 6 August, 10: 228. Isle of 

Oleron, ^S'estern France. 
1854. Heliophoca allantica Gray, Ann. Mag. N.H. /j.- 202. Deserta Grande Island, 

Madeira group. 

Subfamily Cystophorinae 

Genus CYSTOPHORA Nilsson, 1820 

1820. Cystophora Nilsson, Skand. Fauna, Dagg. Djur. /.• 382. Cvstophora borealis 

Nilsson = Phoca cristata Erxleben. 
1826. Stemmatopus F. Cuvier, Diet. Sci. Nat. ^g: 550. Stemmatopus cristatus Cuvier = 

Phoca cristata Erxleben. 
191 1. Cystophoca Brass, Aus dem Reiche der Pelze, 668. Renaming oi Cystophora. 

I species: Cystophora cristata, page 333 

Cystophora cristata Erxleben, 1777 Hooded Seal , Bladdernose) 

Approximate distribution ol" species: .\rctic Europe, .\sia and North America (see 
Anderson, 1947, Canadian Recent Mammals, 80, for Nearctic range). ". . . the deep part 
of the North-\Vestern Atlantic where it is commonest) and adjoining areas of the 
Arctic Ocean, i.e. it extends from Newfoundland, Labrador and Greenland to 
Spitzbergen and Bear Island, east of which — in the shallower part of Barents Sea — it 
only occurs in certain years and in small numbers. Separate individuals, however, 
sometimes swim great distances: one specimen was caught in the Yenesei, near 
Yeneseisk" (Bobrinskii). Has been recorded also from Norway, France, British Isles, 
Portugal I'Santos, 1936), and during migrations to Danish Straits. 



Cystophora cristata Erxleben, 1777 

1777. Phoca cristata Erxleben, Syst. Regn. Aiiim. /; 590. Southern Greenland and 

1820. Cystophora borealis Xilsson, Skand. Fauna. Dagg. Djur. /.• 383. Locality as 

above, based on Gmelin, 1788 cristata, and in turn Erxleben, 1777. 
1823. Phoca mitrata G. Cuvier, Oss. Foss. 5.' 210. 

FAMILY': Procaviidae, page 334 


Genus: Procaria, page 334 

On this family, see particularly Hahn, 1934, Die Familic der Procaviidae, ^. 
Sauget. g: 207-358. Flower and Lydekker recognized two genera in this family, 
Procavia and Dcndrohyrax, characterized by differences in dentition. Although some 
authors refer all Hyraxes to one genus Procavia, there is considerable evidence in the 
material examined that Dcndrohyrax is valid. It has brachyodont cheekteeth, and in 
fully adult skulls the three upper molars are normally a little shorter than, or sub- 
equal to, the four premolars. Procavia has hypsodont cheekteeth, and in fully adult 
skulls the three upper molars are normally clearly longer than the four premolars. 
Hahn and other authors recognize a third genus, Heterohyrax, which does not differ 
from Dcndrohyrax in dentition, but which has the orbit not ringed by bone, whereas 
Dcndrohyrax usually has it ringed by bone. But as the character is not strictly constant 
in South African Dcndrohyrax, it is difficult to see how Heterohyrax could be more than a 
subgenus of Dcndrohyrax. Hahn retained four species in Procavia, two of which, 
habessinica and rujiceps, are supposed to occur in the Palaearctic region. He gives very 
little evidence that these two species are in reality morphologically definable when 
compared with the earliest named Procavia capensis from the Cape. One of us 
(T. C. S. ^L-S.) has not found his characters of the first lower premolar constant in 
habessinica races; his measurements of the skulls and teeth for the three species over- 
lap; and until the contrary is proved, we prefer to regard both the supposed northern 
species as further races of P. capensis. 

Genus PROCAVIA Storr, 1780 

1780. Procavia Storr, Prodr. Meth. Mamm. 40, pi. B. Cavia capensis Pallas. 
1783. Hyrax Hermann, Tabl. Affin. Anim. 115. Cavia capensis Pallas. 
1868. Euhyrax Gray, Ann. Mag. N.H. /.• 46. Hyrax hahessintcus Hcmprich & Ehren- 

I species in the area covered by this list: 
Procavia capensis, page 335 



Procavia capensis Pallas, 1766 Hyrax, "Cony" or Dassie 

Approximate distribution of species (as here understood) : Arabia, Palestine, Sinai, 
Syria; Algeria; Libya. From Somaliland, Sudan, Northern Nigeria, Asben and 
Senegal southwards to Cape Town, George and Albany districts in Cape Province, 
where it is very common. 

(Procavia capensis capensis Pallas, 1766. Extralimital) 

1766. Cavia capensis Pallas, Zool. Misc. 30, pi. 3. Cape of Good Hope. 

Procavia capensis syriaca Schreber, 1784 

1784. Hyrax syriacus Schreber, Saugeth. pi. 240B: 1792, 4: 923. Mt. Lebanon, 

Syria. (See Moreau, Hopkins & Hayman, 1946, P.Z.S. ii§: 431.) 
1868. Hyrax sinaitkus Gray, Ann. Mag. N.H. /.■ 45. Locality not given; probably Mt. 

Sinai, Sinai Peninsula. 
1917. Procavia sinailica ehrenbergi Brauer, S.B. Ges. Nat. Fr. Berlin, 301. El Tor, near 

\Vadi Timar, Sinai. 
19 1 7. Procavia sinailica schmidtzi Brauer, loc. cit. 302. Mountain of Bteha Plain, north 

of Lake Galilee, Palestine. 

Hahn (1934) restricted syriacus Schreber to Abyssinia, for the same reason that 
Gray had done, namely because Schreber quotes largely from Bruce in describing 
this hyrax and also having regard to their interpretation of Schreber's plate. But we 
agree with Thomas (1892) that Schreber clearly intended the Syrian form as well as 
the Abyssinian form, and that both from the text and from the title " Der syrische 
Klippschliefer" there is every ground for including the Syrian form under syriacus 
rather than excluding it, and we agree that Mt. Lebanon was rightly selected as the 
type locality. As no member of the subgenus Heterohyrax is known to occur in Asia, the 
earliest name for that wholly African group will be Hyrax brucei Gray, 1868, from 
Abyssinia. The type oi Heterohyrax should be quoted as Dendrohyrax blainvillii Gray = 
Hyrax brucei Gray. 

Procavia capensis burtoni Gray, 1868 

1868. Hyrax burtonii Gray, Ann. Mag. N.H. /; 43. "Egypt." Probably extralimital 
(Sudanese) but might occur in extreme Southern Egypt. 

Procavia capensis jayakari Thomas, 1892 

1892. Procavia syriaca jayakari Thomas, P.Z.S. 63. Dofar, Southern Arabia. 

Procavia capensis bounhioli Kollman, 191 2 

1912. Procavia bounhioli Kollman, Bull. Mus. H.N. Paris, 18: 281. Ahaggar, Sahara 

Desert, Algeria. 
1932. Procavia {Heterohyrax) antineae Heim de Balsac & Begouen, Bull. Mus. H.N. 

Paris, 2, 4: 479. Ahaggar, Algeria. 

(G. Allen (1939) follows Hahn in listing the last form as a synonym oi bounhioli on 
p. 451, but lists it as a distinct species oi Heterohyrax on p. 445.) 




FAMII.V: Elcphanlidae, page 336 


Genus: Elcphas, page 336 

Genus ELEPHAS Einnacus, 1758 
1758. Elephas Linnaeus, Syst. Nat. loth ed. /; 33. Elephas maximui Linnaeus. 
I species: Elephas maximia, page 33G 

Elephas tnaximus Linnaeus, 1758 Indian Elephant 

Approximate distributicin of species: Ceylon, India (range modified by human 
agency and domestication); Blanford (i8gi) stated that elephants occurred wild 
along the base of the Himalayas as lar west as Dehra Dun and in places in the great 
forest country between the Ganges and Kistna, in the Western Ghats and Mysore. 
Assam, Burma, Siam, Cochin-China. Malay States, .Sumatra. (Introduced in Borneo. 
Deraniyagala, 1950, Proc. ^th Ann. Session Ceylon Assoc. Sci. 10, quotes Laufer (1925) 
as evidence for the elephant being certainly indigenous in Bcirnco, but an examina- 
tion of Laufer does not bear this out.) 

On the races, see Pocock, 1943, .1"". Mag. .N'.H. 10: 273, and Chasen, 1940, 
Handlist Malaysian Mammals, 190 (footnote). 

Elephas maximus maximus Linnaeus, 1758 

1758. Elephas maximus Linnaeus, Syst. Nat. loth ed. /.• 33. Ceylon. 

i()40. Elephas maximus vilaliya Deraniyagala, J. Roy. Asiat. Soc. Ceylon Branch, jj^, 

91: 130, fig. I, 6. Manampitiya, in the flood plain of Mahavili River, 

Eastern Ceylon. Status^(/f- Pocock. 

ElEPH.\S M.XXIMLS INDK.IS G. Cuvicr, I 797 

17117. Elejihas iniiicus Cu\ier, Tabl. Elem. H.N. 148. Ceylon. Slierborn dates indicus 
Cuvier from Mem. Inst. Paris, i',- 21, of 1798 (27 September), but this is 
antedated hy indicus Cu\ier, Tabl. Elem. H.N. 148, which was noticed on 
24 December 171)7, and therefoic published some time before that date. 

(?) 1797. Elephas aualicus Blumeiih.K li, Hand. Naturg. ed. 5, 124. "Asia, chiefly 

184 -). Elephas nidicui liengahnsis Blain\ille, Osteogr. .M.imm. 3-,;;, pi. iii. Bengal. 

1916. F.leptuts mnxmnn maximus of L\(lckker, Lngulates B.M. 5.' 82; not ol 
LiniKiciis, I 7-,;!. 

Range: the mainl.ind i-.mge of the species. Pcxdck i alK the mainland elephants 
A", m. hengalensis .md it is not clear why he discards the rarlii 1 n.ime indicus. 

Elephas maximius cievianicis Blaiii\ill<-, 1845 

1845. Elephas 1)1, Inns cevhinieiis Blaiinillc, Osteogr. .Nfuniii. 3-1",, pi. iii. Ceylon. 


FAMILY: Dugongidae, page 337 
There are two living families, but only one of them comes into our region. 

See Pocock, 1940, Some Notes on the Dugong, Ann. Mag. M.H. 5; 329. 
Genus: Dugong, page 337 

Genus DUGONG Lacepede, 1799 

1799. Dugong Lacepede, Tabl. Mamm. 17. Dugong indicus Lacepede. 

1803. Platystomus Fischer, Nat. Mus. Paris, 2: 353. Platjstomus dugong Gmelin = 

Trichechus dugon Muller. Not Platystoma Meigen, 1803, an insect. 
1808. Dugungus Tiedemann, Zoologie, /.• 554. Emendation. 
181 1. Halicore Illiger, Prodr. Syst. Mamm. et Avium, 140. Trichechus dugong Gmelin 

= Trichechus dugon Miiller. 
182 1. Dugongidus Gray, London Med. Repos. /j; 309. Trichechus dugon Muller. 

I species: Dugong dugon, page 337 

Dugong dugon Muller, 1776 Dugong 

Approximate distribution of species: has been recorded from seas of Portuguese 
East Africa, Madagascar, Mafia Island (off Tanganyika), Kenya, the Red Sea, 
coasts of Malabar, India, Ceylon, the Andaman Islands and Mergui Archipelago, 
Liukiu Is., Formosa, Malaysian Seas, Philippine Islands, and to Northern Australia. 
Doubtless exterminated in some of these places. 

Dugong dugon Miiller, 1776 

1776. Trichecus (sic) dugon Muller, Linne's Vollstandingen Natursyst. Suppl. 21. 

Cape of Good Hope to the Philippines. 

1777. Trichechus dugung Erxleben, Syst. Regn. Anim. 599. 

1799. Dugong indicus Lacepede, Tabl. Mamm. 17. Indian Ocean. 
181 1. Halicore dugong Illiger, Prodr. Syst. Mamm. et Avium, 141. 
1833. Halicore hemprichii Ehrenberg, in Hemprich & Ehrenberg, Symb. Phys. Mamm. 
2: folio k (footnote). Barkan Island, Red Sea. 

1833. Halicore lottum Ehrenberg, in Hemprich & Ehrenberg, loc. cit. Hauakal Island, 

southern part of Red Sea. 

1834. Halicore tabernaculi Ruppell, Mus. Senckenburgianum, /; 113, pi. 6. Red Sea 

(based on a skeleton found on Maxud Island). 
1877. Halicore cetacea Heuglin, Reise in Nordost. Afr. 2: 135. Red Sea. 

The name hemprichii is available if the Red Sea race can be proved distinct from 
that of the Indian Ocean. G. Allen lists it as a synonym oi dugon; but Pocock, (1940, 
330) does not feel justified in adding hemprichi definitely to the synonymy oi dugon. 




FAMILIES: Equidae, page 340 

Rhinocerotidae, page 339 
Tapiridae, page 338 

This is a relict order, with many fossil families and genera but only a handful of 
surviving species. Simpson (1945) divided the existing Perissodactyla into two sub- 
orders, the Hippomorpha for the Ecjuidae, and the Gcratomorpha for the Tapiridae 
and Rhinocerotidae. Each of the last-named families is the type of a distinct super- 
family. Blanford, 1891, Fauna of British India, 468-479, gives short summaries of the 
main diflercnces between the families and most of the species of Asiatic Perissodactyla. 



Genus: Tapirus, page 338 

Genus TAPIRUS Brisson, 1762 

1762. Tapirus Brisson, Regn. Anim. 81-82. Tapirus terrestris Brisson = Hippopotamus 
terrcstris Linnaeus, from Brazil. Hopwood, 1947, P.Z.S. 1 ly, 533-536, would 
disregard Brisson and date Tapirus IVom Brunnich, 1771, Zool. Fundamenta, 
44, 45, with type Hippopotamus terrestris Linnaeus. 

1779. Tapir Blumenbach, Handbuch Naturg. /.' 129. 

1830. Rfiinoc/werus VVagler, Syst. Nat. Amphib. 17. Substitute for Tapirus Brisson. 

1872. Tapvra Liais, Climats Geol. 397. Emendation oi' Tapirus. 

1 91 3. Acrocodia Goldman, Proc. I5iol. Soc. Washington, 26: 65. Tapirus indicus 
Desmarcst. Valid as a subgenus. 

I species in Asia: 

Tapirus indicus, page 338 

We follow Simpson in referring all living tapirs to one genus, but differences in 
the cranium, and in the colour pattern of the adults, seem to justify subgeneric 
distinction between the Asiatic species and its South American allies. 

Subgenus ACROCODLi Goldman, 191 3 

Tapirus indicus Desmarest, 181 9 Malayan Tapir 

Approximate distribution of species: Sumatra and Malay Peninsula, as far north 
as the Burmo-Siamcse borders in latitude 18° N. 

Tapirus indicus Desmarest, 1819 

1 81 9. Tapirus indicus Desmarest, Nouv. Diet. H.N. J2: 458. Malay Peninsula. Range: 
as above. 




Genera: Didermocerus, page 340 
Rhinoceros, page 339 

The prior generic name for the Asiatic Two-horned Rhinoceros is Didermocerus 
Brookes, 1828. Simpson (1945) calls this Dicerorhinus, and suggests, somewhat half- 
heartedly, that the name Didermocerus may conveniently be dropped, on the ground 
of its publication in a sale catalogue. This in itself is no bar to "publication" within 
the meaning of the Regies, and the catalogue was on sale to the public for half a 
crown. Moreover, Simpson adopts Acinonyx, which appears in the same publication. 

Simpson (1945) lists the living rhinoceroses in two subfamilies: the "Dicero- 
rhininae" with Dicerorhinus {= Didermocerus), Ceratotherium and Diceros; and the 
Rhinocerotinae with Rhinoceros. But this arrangement, as Pocock, 1945, P-Z-S- 114: 
437, points out, gives undue importance to the possession of two horns or one, and 
obscures the fact that in cranial and dental characters the Asiatic rhinoceroses clearly 
form one group and the African ones another. We therefore follow Pocock in dividing 
the living rhinoceroses into the Rhinocerinae (or Rhinocerotinae, as the word should 
have been formed) with Rhinoceros and Didermocerus, and the Dicerinae (or Dicero- 
tinae) with Diceros and Ceratotherium. Pocock (1945, 449) gives a key based on this 

Subfamily Rhinocerotinae 

Genus RHINOCEROS Linnaeus, 1758 

1758. Rhinoceros Linnaeus, Syst. Nat. loth ed. /.• 56. Rhinoceros unicornis Linnaeus. 
1867. Eurhinoceros Gray, P.Z.S. 1009. Rhinoceros unicornis Linnaeus. 

2 species : Rhinoceros sondaicus, page 340 
Rhinoceros unicornis, page 339 

For key to these species, see Blanford (1891, 472). 

Rhinoceros imlcornis Linnaeus, 1758 Great One-horned Rhinoceros 

Approximate distribution of species: Nepal, Bihar, Bengal Duars, Cooch Behar, 
Assam. Becoming rare. 

Rhinoceros unicornis Linnaeus, 1758 

1758. Rhinoceros unicornis Linnaeus, Syst. Nat. loth ed. /.• 56. Probably the sub- 
Himalayan Terai of Assam (Lydekker). 

1817. Rhinoceros indicus Cuvier, Regn. Anim. /; 239. 

1830. Rhinoceros asiaticus Blumenbach, Hand. Naturg. ed. 12, 107. No locality 

1867. Rhinoceros stenocephalus Gray, P.Z.S. 1018. Asia. 


PALAEARCrnt; and INDIAN MAMMALS 1758-1946 

Rhinoceros sondaicus Desmarest, 1822 Lesser One-horned Rhinoceros 

Approximate distribution of species: Burma, Siam, Cochin-China, Malay States, 
Sumatra, Java. Now a rare animal. 

Rhinoceros sond.mcus Desmarest, 1822 

1822. Rhinoceros sondaicus Desmarest, Mammalogie, 2: 399. Java. 

1840. Rhinoceros inermis Lesson, Compl. de Buflbn, /.■ 514. Sunderbans, mouths of 

the Gan,s;es, India, nom. nud.,fide Sherborn. 
1 868. Rhinoceros foiveri Gray, V.Z.i. 1867; loi 5. Sumatra (not in Chasen's list, 1940). 

.See Loch, 1937, The Ja\an or Lesser One-horned Rhinoceros and its geographical 
distribution, J. Malayan Branch R. Asiat. Soc. /j, 2: 130. 

Genus DIDERMOCERUS Brookes, 1828 

1828. Didermocerus Brookes, Cat. Anat. Zool. Museum of J. Brookes, London, 75. 
Diderrnocerus sumalrensis = Rhinoceros sumalrensis Fischer. 

1 84 1. Diccrorhinus Gloger, Handbuch Naturgesch. 125. Rhinoceros sumalrensis Cuvier. 
1868. Cerntnrhinus Gray, P.Z.S. 1867: 1021. Rhinoceros sumalrensis Guvier. 

I species: Didermoceru\ sumalrensis, page 340 

Didermocerus sumatrensis Fischer, 1814 Asiatic Two-horned Rhinoceros 

Approximate distribution of species: ? Assam, Burma, Siam, ? Indo-China, Malay 
States, Sumatra, Borneo. Becoming a rare animal. 

(Didermocerus sumatrensis sumatrensis Fischer, 1814. Extralimital) 
1814. Rhinoceros sumalrensis Fischer, Zoogn. j: 301. Sumatra. Range: Sumatra and 

Didermocerus sumatrensis lasiotis Buckland, 1872 

1872. Rhinoceros lasiolis Buckland, Land and \Vater, 10 August. See Harper, 1940, 

J. Mammal. 21: 201. South of Chittagong, Eastern Bengal. 
(?) 1854. Rhinoceros crossii Gray, P.Z.S. 251. Locality unknown. (Based on a horn 
which could equally well have come from an African rhinoceros.) 

1873. Ceralorhinus niger Gray, Ann. Mag. N.H. //.• 357, pi. 11. Malacca. Not of 

Schinz, 1845. 
1873. Ceralorhinus blvlhii Gray, Ann. NLig. N.H. //.• 3G0. Tenasserim. 
Range: ? Assam, Burma, .Siam, ? Indo-China, Malay States. 



Genus: Equus, page 341 


Genus EQUUS Linnaeus, 1758 

1758. is^i/w Linnaeus, Syst. Nat. lothed. /.■ y^. Equus caballus hinnaeus, the domestic 

1762. Asinus Brisson, Regn. Anim. 70. Equus asinus Linnaeus. Valid as a subgenus. 
1762. Onager Brisson, Regn. Anim. 72. Equus asinus Linnaeus. 
1824. Asinus Gray, Zool. Journ. /.• 244. Equus asinus Linnaeus. 
1924. Mitroluppus Matschie, S.B. Gcs. Nat. Fr. Berlin, n)22: 68. Microhippus lafeli 

Matschie = Equus kiang Moorcroft. 

Simpson also quotes a name, Hemionus Cuvier, 1823. The only reference we have so 
far traced is Hemionus Cuvier, 1821, Diet. Sci. Nat. 555, which seems to be a trivial, 
not a generic name. 

There are other, extralimital (African) subgeneric names. 

For the geographical distribution of recent Equidae see Antonius, 1938, P.^.S. 
107B: 557. 

2 species in Asia: 

Equus hemionus, page 341 
Equus przewalskii, page '^/^i 

For key to these species, see G. Allen, 1940, Mammals of China and Mongolia, 2: 
1 28 1. Bobrinskii (1944) refers hemionus to the subgenus Asinus, but this is more 
usually restricted to Equus asinus Linnaeus which now occurs as a wild animal only 
in Eastern Africa (Sudan, Somaliland). 

Equus przewalskii Poliakov, 1881 Przewalski's Horse, Tarpan 

Approximate distribution of species: Mongolia, Chinese Turkestan. 

Equus PRZEWALSKII Poliakov, 1 88 1 

1 88 1. Equus przewalskii Poliakov, Proc. Imp. Russian Geogr. Soc. ly, i : pis. i and 2. 
See also 1881, Ann. Mag. N.H. 8: 16. Oasis of Gashun (44°30' N., 90° E.), 
steppe country of Eastern Zungaria. (Harper, 1940, J. Mammal, 21: 196.) 

1903. Equus hagenbecki Matschie, Naturwiss. Wochenschrift, 18, 49: 583. Ebi Spring, 
Gobi Desert, Mongolia. 

Lydekker considered this to be a subspecies of Equus caballus Linnaeus, the 
domestic Horse. 

Equus hemionus Pallas, 1775 Asiatic Wild Ass 

Approximate distribution of species: Southern and Eastern Russian Turkestan 
(now surviving round Kushka, and in the south-west of the Balkash basin; single 
arrivals from China have recently occurred in frontier area of Hi Valley (Bobrinskii) ), 
Mongolia, Chinese Turkestan, Tibet; Persia, Iraq, Syria; Ladak, Baluchistan, Nepal, 
Sind and Cutch; Afghanistan. 

For a discussion of the type localities, status, etc. of these Asses, see Harper, 1940, 
J. Mammal. 21: 197; also Pocock, 1948, P.^.S. iiy: 764. 



Eqi'L'S HEMiONUS HEMiONUs Pallas, 1 775 Chigetai, Kulan, or Mongolian Wild Ass 

1775. Equus hemionus Pallas, Nov. Comm. Ac. Sci. Petrop. ig: 394, pi. 7. Tarei- 

Nor, Dauria, Transbaikalia (50° N., ii5°E.). 
1891. Equus hemionus var. typtcus Sclater, Cat. Mamm. Ind. Mus. 2: 198. 
(?) 1904. Equus onager castaneus Lydekker, Nov. Zool. //.• 590, pi. .wiii. Kirghis Nor, 

Kobdo, Western Mongolia. 
191 I. Equus (Asinus) hemionus bedfordi Matschie, in Futtcrcr, Durch Asien, j, 5, 

Zoolog. Nachtrag, 23. Probably Kobdo, Mongolia. 
191 1. Equus [Asinus) hemionus luteus Matschie, loc. cit. 24. Western Gobi. 
Range: now apparently only found about Orok Nor and Zagan Nor, in Central 

Equus hemionus onager Boddaert, 1 785 Persian Onager or Ghor-khar 

1785. Equus onager Boddaert, Elench. Anim. 160. Kasbin, North-Western Persia, 

near the Caspian. 
1891. Equus onager var. typicus Sclater, Cat. Mamm. Ind. Mus. 2: 198. 
(?) 191 1. Equus (Asinus) hemionus Jinschi Matschie, in Futterer, Durch Asien, j, 5, 
Zool. Nachtrag, 24. North-east of Zaisan Nor, Scmipalatinsk, Russian Asia. 
Range: north-eastern parts of Persia and North-Western Afghanistan; Russian 
Turkestan, as abo\e. 

Equus hemionus khur Lesson, 1827 Indian Wild Ass or Ghor-khar 

1827. Equus khur Lesson, Mammalogie, 347. The Little Rann of Cutch, India. 
(?) 1841. Asinus hamar H. Smith, Jardines Nat. Libr. Mamm. 31: 351, pi. 19. Pro- 
vince of Pars, Persia, between Yezdi Khast and Shulgastan. 
1862. Asinus indieus Sclater, P.Z.S. 163, nom. nud. 
1869. Equus indieus George, Ann. Sci. Nat. Zool. 12: 35. 
Range: the Rann of Cutch, possibly Baluchistan, and South-Eastern Persia. 

Equus hemionus kiang Moorcroft, 1841 Kiang 

1841. Equus kiang Moorcroft, Travels in the Himalayan Provinces, /.• 312. Eastern 

parts of Ladak, Kashmir. 

1842. Asinus equioidei Hodgson, J. Asiat. Soc. Bengal, //, i : 287. Plains of Tibet. 
1847. Asinus poly don Hodgson, Calcutta J.N. H. j: 469. Hundes district of Tibet. 
1869. Asinus kyang Kinloch, Large Game Shooting in Thibet, /.• 13 Tibet. 

191 1. Equus (Asinus) kiang holdereri Matschie, in Futterer, Durch Asien, j, 5, Zool. 

Nachtrag, 29. South-western shore of Lake Kukunor, Chinese Central Asia. 
1924. Mierohippus lafeli Matschie, S.B. Ges. Nat. Fr. Berlin, i()22: 68. Tosson Nor, 

Range: Ladak, Nepal, Sikkim, Tibet to Kukunor district. 

Equus hemionus hemippus I. Geoffroy, 1855 

1855. Equus hemippus I. Geoffroy, C.R. Ac. Sci. Paris, 41: 1214, 1220. Syria. 
1869. Equus hemionus var. syriacus Milne-Edwards, Nouv. Arch. Mus. H.N. Paris, 5, 

Bull.: 40, pi. 4. Damascus, Syria. 
Range: Syrian Desert and adjacent parts. Possibly now extinct. 




Responsibility for the classification of this Order is taken by T. C. S. M.-S. 

^Vorks of reference : 

Allen, G. 1939. Checklist African Mammals, Bull. Mus. Comp. ^ool. Harvard, 83. 

1 940. Mammals of China and Mongolia, 2. 

BoBRiNSKii, KuzNETZOV & KuzYAKiN. 1 944. Mammals of the U.S.S.R. 
Lydekker. 1 91 3-15. Catalogue of Ungulate Mammals in the British Museum, i-^. 

i8g8. The Deer of all Lands. 

Miller. 1912. Catalogue of Mammals of Western Europe. 

PococK. 191 1. On the specialized cutaneous glands of Ruminants, P.^.S. igio: 

1 91 8. On some external characters of ruminant Artiodactyla, Ann. Mag. N.H. 

i: 426-435; 2: 125-144, 214-225, 367-374, 440-459. 

1919- On the external characters of existing Chevrotains, /'.,<]'. i?. I. 

1923. On the external characters of Elaphurus, Hydropotes, Pudu and other 

Cervidae, P.Z-S. 181. 

1923. External characters of the Pygmy Hippopotamus, and of the Suidae and 

Camelidae, P.Z-S. 531- 

ScHWARZ. 1937. Wiss. Ergeb. Oldowaj-Exp. igij, N.S. pt. 4; 7-90, 3 pis. 

ScLATER & Thomas. 1894-1900. The Book of Antelopes, 1-4. 

Simpson. 1945. Principles of Classification and Classification of Mammals, Bull. 

Amer. Mus. N.H. 8§. 
WiNGE. 1924. Pattedyr-Slaegter, j. 

Although his keys and specific diagnoses are not always very clear, Lydekker's 
Catalogue of Ungulate Alammals is one of the most useful works on this Order. Simpson 
(1945) classified the living Artiodactyla of the Palaearctic and Indian regions as 
follows : 

Suborder: SUIFORMES 
Infraorder: Suina 
Family: Suidae 

Suborder: TYLOPODA 
Family: Camelidae 

Infraorder: Tragulina 
Family: Tragulidae 

Infraorder: Pecora 

Superfamily: Cer\'oidea 

Family: Cervidae 
Superfamily: Bovoidea 

Family: Bovidae 


PAI.AF,ARCTK: and INMMAM mammals 1 758-1946 

This arrangement is in general agreement with most of the earlier authors and is 
here followed. Subfamilies will be discussed below in the appropriate places. 
FAMILIES: Bovidae, page 377 

Camelidae, page 348 
Cervidae, page 352 
Suidae, page 344 
Tragulidae, page 349 

The Hippopotamidae (genus Hippopotamus) inhabited the Lower Nile Valley till 
about 1 81 5, but are no longer found in the Palaearctic region. 



Genus: Sus, page 344 

Genus SUS Linnaeus, 1 758 

1758. Sus Linnaeus, Syst. Nat. loth ed. /.• 49. Sus scrofa Linnaeus. (Opinion 75 of the 
International Commission on Zoological Nomenclature.) 

1847. Porciila Hodgson, J. Asiat. Soc. Bengal, 16: 423. Porcula salvania Hodgson. Valid 
as a subgenus. 

1862. Centuriosus Gray, P.Z.S. 17. Sus plicireps Gray (a Japanese domestic \-ariety). 

1868. Scrofa Gray, P.Z.S. 38. Domestic Pig. [Sus domrstkus Brisson = Sus scrofa 


1869. Euhys Gray, Gat. Carnivora, etc. Brit. Mus. 339. Sus barbatus Miiller, from 

1873. Aulacochoerus Gray, Ann. Mag. N.H. //.• 435. Sus vittalus Miiller = Sus vittalus 

Boie, from Sumatra. 
1873. Dasvchoerus Gray, Ann. Mag. N.H. //.• 435. Sus verrucosus Muller & Schlegel, 

from Java. 
1892. Sinisus Heude, Mem. H.N. Emp. Ghinois, 2: 102. Apparently based on the 

Chinese forms of Sus scrofa. 

■2 species in the area covered by this list: 
Sus salvanius, page 348 
Sus scroja, page 345 
.S>/.* Milranius is separated subgenerically as Porcula on account of its small size, the 
\ery short tail, and there being only three pairs of teats as opposed to six pairs in Sus. 
The other wild pigs of the region are here treated as belonging to a single species, 
.S>/( Hrofa. It may be as well to draw attention to the fact that Chasen, 1940, Handlist 
of Malaysian Mammals, besides the species >S'. verrucosus and .S'. barbatus recognized one 
species of wild pig in the Malaysian region, which he listed as Sus crrstalui with 
vittalus as a race. But he should have done it the other way round, since he correctly 
referred vittalus to Boie, 1828, Pjijdr. .Nat. W'etcnsch. 3, i : 240, which antedates crislatus 
by ele\en years. Both eristatus and viltatw. are here regarded as representing S. scrnja. 

Subgenus SUS Linnaeus, ry^S 

Sus scrofa Linnaeus, 1758 Wild Boar 

Approximate distribution of species: Continental Europe, known from Spain and 
Portugal, France, Belgium (Holland and Denmark, became extinct but reintroduced 
after 1800), Germany, Switzerland, Italy, Corsica and Sardinia, Baltic States (south 
of 58°N.), Poland, Czechoslovakia, Austria, Hungary, Yugoslavia, Rumania, Bul- 
garia, Greece. In Western Russia, roughly from Riga towards Velikie Luki, but 
turning south before reaching there, passing round west of Vitebsk and roughly 
along the White Russian frontier, Chernigov district included, to Kiev, and a little 
south of Mogilev, reaching the Dniester, which it follows to the Black Sea (with 
individual cases of incursions fairly far east of this line) (Bobrinskii). Caucasus. 
Widely distributed in Russian Turkestan, and to as far north as Pavlodar on Irtish 
River. Far East of Siberia from eastern Sayan Mountains, through Transbaikalia 
and Amur regions to Ussuri region. Japan, Formosa, Manchuria; Mongolia, Chinese 
Turkestan; all the larger states of China (perhaps excepting Yunnan). Asia Minor, 
Persia, Afghanistan, Palestine. India, from Baluchistan, Kashmir, Nepal south- 
wards through the Peninsula to Ceylon, east to Burma. Indo-China, Siam, Malay 
States, Sumatra, Java and various small islands, Flores. Rio de Oro, Morocco, 
Algeria, the Sudan, and formerly Egypt where it became extinct about 1900 (Flower, 

Sus SCROFA SCROFA Linnaeus, 1758 

1758. Sus scrofa Linnaeus, Syst. Nat. loth ed. /.• 49. Germany. 

1785. Sus setosus Boddaert, Elench. Anim. /.• 157. Substitute for scrofa. 

1 785. [Sus setosus) aper Boddaert, loc. cit. 

1788. Sus scrofa ferus Gmelin, Linn. Syst. Nat. /.• 217. 

181 1. Sus europaeus Pallas, Ross. Asiat. /.• 265. Substitute for scrofa. 

1836. Sus scropha ]-a.rAint, Nat. Libr. Mamm. 5.- 205. Substitute for scrofa. 

1882. Sus scrofa var. celtica Strobel, Atti Soc. Ital. Sci. Nat. Milano, 25.- 79. 

Range: from France and Germany eastwards into Western White Russia. 

Sus SCROFA CRiSTATUS Wagner, 1839 

1839. '^"^ cristatus Wagner, Munch. Gelehrt. Anz. g: 435 (misprinted as "535"). 

Probably the Malabar coast, India. 
1842. Sus aper var. aipomus Hodgson, J. Asiat. Soc. Bengal, 10: 911. Nepal. 

1842. Sus aper var. tsonotus Hodgson, loc. cit. Nepal. 

1843. Sus indicus Gray, List. Mamm. B.M. 185. 

1847. Sus affinis Gray, Cat. Osteol. B.M. 71. Nilgiri Hills, India. 
1851. Sus zeylonensis Blyth, J. Asiat. Soc. Bengal, 20: 173. Ceylon. 
i860. Sus bengalensis Blyth, J. Asiat. Soc. Bengal, 2g: 105. Bengal. 
1900. Sus cristatus typicus Lydekker, Great & .Small Game India, 261. 
Range: Ceylon and Indian range of species above. 



Sus scROFA LEUCOMYSTAX TeiTiminck, 1842 

1842. Sus leucomvstax Tcmminck, Siebolds Fauna Japon. Mamm. 6. Japan. 
1885. Sus vittatus japomca Nehring, Zool. Garten, 26: 336. 
Range includes Islands of Hondo, Shikoku, Kiushiu, Japan. 


1858. Sus andamanensis Blyth, J. Asiat. Soc. Bengal, 2j: 267. Port Blair, Andaman 
Islands, Bay of Bengal. 

Sus scROF.A. BARBARUS Sclater, i860 

i860. Sus saofa var. harharus Sclater, P.Z.S. 443. North Africa. 

1867. Sus scrofa var. alajra Loche, E.xpl. Sci. dc PAlgcrie, Zool. Mamm. 59. Country 

of Beni Sliman, Algeria. 
(?) 1937. Sus scrofa sahariensii Heim de Balsac, Bull. Soc. Zool. France, G2: 333. 

Jebel Guettar, north-west of Ain Sefra, Northern Algeria. 
Range: Morocco, Rio de Oro, Algeria. 

Sus SCROFA TAIVANUS SwiuhoC, 1 863 

1863. Ponula taivana Swinhoe, P.Z.S. 1862: 360. Formosa. 

Sus SCROFA LiBYCUS Gray, 1868 

1868. Sus libycus Gray, P.Z.S. 31. Xanthus, near GUnek, South-Western Asia Minor. 

Sus scROF,*! MOUPiNENsis Milnc-Edwards, 1872 

1872. Sus moupinensis Milne-Edwards, Nouv. Arch. Mus. H.N. Paris, 7, Bull.: 93 

(footnote). Moupin, Szechuan, China. 
1888. Sus oxjodontus Heude, Mem. H.N. Emp. Chin. 2: 54, mm. nud. Upper Han 

River, Shensi, China. 
1888. Sus dicrurus Heude, loc. cit. 55. Divide between Han and Kincha Rivers, 

Shensi, China. 
1892. Sus curtidens, Sus laticeps, Sus collinus and Sui acrocraniui Heude, Mem. H.N. 

Emp. Chin. 2: 1 14. 
1899. Sus planiceps Heude, Mem. H.N. Emp. Chin. 4: 132. Ho Shan, Anhwei, Clhina. 
Range: Szechuan, eastwards to Chihli, Northern China. 

Sus SCROFA NiGRiPES Blanford, 1875 

1875. Sus scrofa var. ruaripes Blanford, J. Asiat. Soc. Bengal, 44, 2: 112. Kashgar 

district, Chinese Tianshan. Ranges throughout Russian Turkestan, and 

probably Afghanistan. 

Sus SCROFA MERiDioN'ALis Forsyth Major, 1882 

1882. Sus scrofa nuridionalis Forsyth Major, Atti Soc. Tosc. Sci. Nat. Pisa, Proc. 

\'erb. 3: 119 (May). Sardinia. 
1882. Sus scrofa var! sardous Strobel, Atti Soc. Ital. Sci. Nat. Milano, 2j: 221 

(September). Sardinia. 



Sus SCROFA ussuRicus Heude, 1888 

1888. Sus ussuricus Heude, Mem. H.N. Emp. Chin. 2: 54. Ussuri Valley, Eastern 


1889. Sus leticomystax var. continentalis Nehring, S.B. Ges. Nat. Freunde Berlin, 141. 

Vladivostock, Eastern Siberia. 
1892. Sus gigas Heude, Mem. H.N. Emp. Chin, 2: 114. Vladivostock, Eastern 

1897. Sus songaricus Heude, Mem. H.N. Emp. Chin, j.- 191. Valley of Sungari River, 

(?) 1897. Sus canescens Heude, loc. cit. 192. Pekin, China. 
1897. Sus mandchuricus Heude, loc. cit. 192. Mukden, Manchuria. 

Sus SCROFA CHiRODONTUs Hcudc, 1 888 

1888. Sus chirodontus Heude, Mem. H.N. Emp. Chin. 2: 54. Poyang Lake, Kiangsi, 

Southern China. 
1888. Sus palustris Heude, loc. cit. (footnote). Not of Riitimeyer, 1861. Valley of 

Yangtze, China. 
1892. Sus leucorhinus, Sus paludosus, Sus melas Heude, Mem. H.N. Emp. Chin. 2: 1 14. 
1899. Sus flavescens Heude, Mem. H.N. Emp. Chin. 4: 130. Yangtze and Taihu, 

Kiangsu, China. 
1899. Sus chirodonticus Heude, loc. cit. Poyang Lake, Kiangsi, China. 
Range: Southern China and Hainan. 

Sus SCROFA coREANUS Heude, 1897 

1897. Sus coreanus Heude, Mem. H.N. Emp. Chin. 5.- 191. Fusan, Korea. 

Sus SCROFA NicoBARicus Miller, 1902 

1902. Sus nicobaricus Miller, Proc U.S. Nat. Mus. 24: 755. Great Nicobar Island, 
Bay of Bengal. 

Sus SCROFA juBATUS Miller, 1906 

1906. Sus jubatus Miller, Proc. U.S. Nat. Mus. jo.- 745. Trang, Lower Siam. Ranges 
north into Indo-China. 

Sus SCROFA ATTiLA Thomas, 191 2 

191 2. Sus attila Thomas, Abstr. P.Z.S. 13; P.Z.S. 393. Kolozsvar, Transylvania. 
Ranges eastwards to the Caucasus and. Northern Persia. 

Sus SCROFA CASTiLiANUs Thomas, 1912 

191 2. Sus scrofa castilianus Thomas, Abstr. P.Z.S. 13; P.Z.S. 392. Quintanar de la 

Sierra, near Burgos, Northern Spain. 
(?) 1912. Sus scrofa baeticus Thomas, Abstr. P.Z.S. 14; P.Z.S. 393. Coto Donana, 

Huelva, Southern Spain. 


1 91 8. Susfalzfeini Matschie, S.B. Ges. Naturf. Fr. Berlin, No. 8, 5. Naliboki, North- 
Eastern Poland. 

2 347 


Sus scROFA RiuKiuANUs Kuroda, 1924 

1924. Sus leucomystax riukiuanus Kuroda, on New Mammals from Riu Kiu Islands 

(Tokyo), II. Kabira, Ishigakijima, Riukiu Islands. 

Sus SCROFA REisFRi Bolkay, 1925 

1925. Sui atlila tfisfii Bolkay, Nov. Mus. Sarajevo, /.■ 13. Bosnia, Yugoslavia. 

Sus sc.ROF.\ MAjoRi dc Beaux & Festa, 1927 

1927. Sus scrofa majori de Beaux & Festa, Mem. Soc. Ital. Sci. Nat. Milano, 9.- 270. 
Mt. Pescali, Tuscany Maremma, Italy. 

Sus SCROFA R.-\DDE.-\Nus Adlerberg, 1930 

1930. Sus scrofa raddeanus Adlerberg, C.R. Acad. Sci. U.R.S.S. 95, figs. 2, 3. Sugu 
Nor, southern Kentai Mountains, Mongolia. Ranges to Southern Trans- 

Subgenus PORCULA Hodgson, 1847 

Sus salvanius Hodgson, 1847 Pygmy Hog 

Approximate distribution: the Terai of Sikkim, Nepal and Bhutan, India. 

Sus S.A.LVANIUS Hodgson, 1847 

1847. Porcula salvania Hodgson, J. Asiat. Soc. Bengal, 16: 423, pis. 12, 13. Sikkim 

Terai, India. 
1863. Sus lilliputensis Gray, Cat. Hodgson's Cloll. B.M., 2nd ed. 15, nom. nud. 

Tate (1947, 311) suggests that salvanius may have been based on young specimens 
of the Indian wild boar. This was not so, and the species is valid. The adult skulls in 
the British Museum have an overall length of only 150-160 mm. 



Genus: Camdus, page 348 

Genus CAMELUS Linnaeus, 1 758 

1758. Camelus Linnaeus, Syst. Nat. loth ed. /.• 65. Camehn bactrianus Linnaeus (see 
Opinion 16 of International Commission on Zoological Nomenclature). 

I species known in the wild state: 

( Ml/hi u\ haclnanus, page 349 



Camelus bactrianus Linnaeus, 1758 Bactrian, or Two-hum ed Camel 

Approximate distribution of species: According to Allen (1940) it is nit possible 
to say whether the camels found in Central Asia are truly wild or are descf nded from 
escaped domesticated stock. Harper (1945) says that the wild Bactrian camel still 
exists in restricted numbers in Chinese Turkestan and in Mongolia. The same 
author quotes Leche (1904) and Lattimore (1929) on the anatomical differences 
between Camelus bactrianus bactrianus and C. b. ferus which support the \iew that the 
latter is a genuinely wild animal, not contaminated to any appreciable extent with 
the blood of the domestic animal. According to Bannikov, 1945, \,ool. J. Moscow, 24: 
200, there are wild Bactrian camels in the Gobi in an area bounded by the parallels 
of 42-45° and by longitudes 96-99°. The only camels in the U.S.S.R. are domestic 
ones (Bobrinskii, 1944). The domestic Bactrian camel has a wide distribution in 
Asia. The Dromedary, or One-humped Camel {Camelus dromedarius Linnaeus), is not 
known in the wild state. 

Camelus bactrianus bactrianus Linnaeus, 1758 (Domestic Bactrian Camel) 
1758. Camelus bactrianus Linnaeus, Syst. Nat. loth ed. /.• 65. "Bactria" = Bokhara. 

Camelus bactrianus ferus Przewalski, 1883 (Wild Bactrian Camel) 

1883. Camelus bactrianus ferus Przewalski, Third Journey into Central Asia, 43. Border 

of the Kum-tagh, east of Lob-Nor and north of the Altyn-tagh, Chinese 

Turkestan (Harper, 1940). 



Genus: Tragulus, page 349 

Genus TRAGULUS Brisson, 1 762 

1762. Tragulus Brisson, Regn. Anim. 65. Tragulus indicus Brisson ?=Cen!us javanicus 

1843. Moschiola Hodgson, Calcutta J. N.H. 4: 292. Tragulus mimenoides Hodgson = 
Tragulus meminna Erxleben. Valid as a subgenus. (Thomas, 1916, Ann. Mao-. 
N.H. 18: 72, says that Moschiola Hodgson is invalid because it was published 
in conjunction with a nomen nudum (i.e. mimenoides). But Thomas was mis- 
taken in thinking this was a nomen nudum. The name mimenoides was published 
in proper form by Hodgson in the previous year (see belowj.i 

Hopwood, 1947, P.Z-S. 117: 534, considers Brisson, 1762, an unavailable work 
and holds, further, that the name Tragulus Pallas, 1779, Spicilegia ^ooloiiica, ij: 27, is 
also unavailable. He proposes dating Tragulus from Boddaert, 1785, Elenchus Anima- 
lium, with type species T. pygmaeus Boddaert = Capra pygmaeus (sic) Linnaeus, 1758. 
This selection of type species is, however, most unfortunate, since Capra pygmaeus 
Linnaeus, 1758, is the Royal Antelope of West Africa, a member of the Bo\-idae. 



An examination of Boddacrt shows, however, that Boddaert should not, by his 
own definition, have included pygmaeus in his Tragt/ltis, since the latter is defined as 
being hornless whereas Moschus pygmaeus Linnaeus, 1766, which is what Boddaert 
definitely quotes, and which is the same thing as Capra pygmaea Linnaeus, 1 758, has 

The only hornless species included by Boddaert in Tragulus are T. moschus 
(= Moschus moschiferus Linnaeus, 1758) and T. mcminna Erxleben, 1777, and in his 
index on p. 49, he writes: "Tragulus {Moschus Linn.)". The type species of Tragulus 
Boddaert, 1785, is therefore Moschus moschiferus Linnaeus, 1758, the Musk Deer. So 
irrespective of Hopwood's curious selection of type species the dating of Tragulus 
from Boddaert, 1785, would be a most unfortunate affair, involving, as it would, the 
transferring of Tragulus away from the Tragulines, with all the resulting confusion. 

Therefore, pending a decision by the International Commission on Zoological 
Nomenclature, we propose to continue dating Tragulus from Brisson, 1762. 

The classification of this genus presents no great difficulties, thanks to the work of 
Lydekker, Chasen and others. Two species of the typical subgenus occur side by side 
more or less from Tenasserim to Borneo. T. meminna, from Western India, is separated 
subgenerically as Moschiola. Pocock gave it generic rank, but we prefer to follow 
Simpson and regard it as a subgenus. See Pocock, iqig, P.^-^- '; ^ind Lydekker, 
191 5, Cat. Ung. Mamrn. B.M. 4: 260, for specific characters. 

3 species : Tragulus javanicus, page 35 1 
Tragulus meminna, page 350 
Tragulus napu, page 351 

Subgenus MOSCHIOLA Hodgson, 1843 

Tragulus meminna Erxleben, 1777 Indian Spotted Chevrotain (Mouse-deer) 
Approximate distribution of species: Ceylon and Peninsular India. (See Cham- 
pion, 1929, J. Bombay N.H. Soc. 23- 985, for Indian details); in India, north 
approximately to Central Provinces. 

Tragulus meminna Erxleben, 1777 

1777. Moschus meminna Erxleben, Syst. Regn. Anim., Mamni. 322. Ceylon. 

(?) 1842. Tragulus mimenoidcs Hodgson, J. Asiat. Soc. Bengal, 10: 914. Nepal Terai. 

1843. Meminna indica Gray, List. Mamm. B.M. i72. Not of Brisson, 1762, but based 

on meminna Erxleben. 
1843. Meminna tnalaccensis Gray, List. Mamm. B.M. 172. Locality unknown. (Gray 

gives "Singapore.") 

Subgenus TRAGULUS Brisson, 1762 

See Kloss, 19 18, J. Fed. Malay Stales Mus. y: 245; Notes on Malayan and other 



A. van Bemmel, 1949, Treubia, 20, 2: 378, points out the necessity for "a rather 
disagreeable change of nomenclature" in this subgenus. For years the Larger 
Mouse-deer has been called Tragulus javanicus Osbeck, and the Lesser Mouse-deer 
T. kanchil Raffles. It was not till 1929 that Sody and Dammerman began to query 
the existence of the Larger Mouse-deer in Java. 

Now van Bemmel has investigated the problem thoroughly and finds that in no 
collection can a specimen of the Larger Mouse-deer be found which is reliably known 
to have come from Java, and, further, that the Larger Mouse-deer has never been 
observed in Java. Furthermore, a study of the description oi javanicus together with 
Osbeck's earlier, pre-Linnean description, and his narrative, makes it clear that the 
animals which Osbeck bought alive from natives on the coast of Udjon Kulon 
Peninsula, Western Java, were Lesser Mouse-deer, which is in fact the only form of 
Mouse-deer which does occur in Java. 

A switch round of names is therefore necessary and the Lesser Mouse-deer must be 
called Tragulus javanicus Osbeck, 1765 (= Moschus kanchil Raffles, 1821), and for the 
Larger Mouse-deer there is Tragulus napu F. Cuvier, 1822 (= Tragulus javanicus 
auct. nee Osbeck) . 

Presumably the form from Java listed by Chasen (1940, 201) as T. kanchil focalinus 
is a synonym of T. j. javanicus, and presumably kanchil Raffles will stand as the 
Sumatran race oi javanicus. 

Tragulus napu F. Cuvier, 1822 Larger Malay Chevrotain (Mouse-deer) 

Approximate distribution of species: Tenasserim, Siam, Indo-China, Malay 
States, Sumatra, Borneo, many small adjacent islands, including Balabac. 

Tragulus napu napu F. Cuvier, 1822 

1822. Moschus napu F. Cuvier, in Geoffroy & Cuvier, H.N. Mamm. 2, 37 : 2. Southern 

Sumatra (Sody, 193 1). 
1900. Tragulus canescens Miller, Proc. Biol. Soc. Washington, ij: 185. Trang, Lower 

Ranges to Tenasserim. 

Tragulus napu versicolor Thomas, 1910 

1910. Tragulus versicolor Thomas, Ann. Mag. N.H. §: 535. Nhatrang, Annam, Indo- 

Tragulus javanicus Osbeck, 1765 Lesser Malay Chevrotain (Mouse-deer) 

Approximate distribution of species: Tenasserim, Indo-China, Siam, Malay 
States, Sumatra, Java, Borneo, and many small adjacent islands. 

(Tragulus javanicus javanicus Osbeck, 1765. Extralimital) 
(?) 1762. Tragulus indicus Brisson, Regn. Anim. 65. (Unavailable, see page 3.) 
1765. Cervus javanicus Osbeck, Reise nach Ostindien und China, 357. Udjon Kulon 
Peninsula, Western Java (van Bemmel, 1949). 


PALAEARC:TIC and IXUIAX mammals 1758-1946 

Tragvlus javanicus affims Gray, 1861 

1861. Traaului ajfinis Grav, P.Z.S. 138. Cambodia, Indo-China. iSee Ossjood, 1932, 

Field Mus. N.H.' Zool. /<?/329.) 
(P"! 1902. Tragulus ravus Miller, Proc. Biol. Soc. Washington, /j; 173. Trang, Lower 

Siam. Ranges to Tenasserim. 
1903. Tragtilns kanchil pierrei Bonhote, Ann. Mag. X.H. //.• 293. Bien Hoa, Lower 

Cochin-China. javanicus lampensis Miller, 1903 

1903. Tragulus lampensis Miller, Proc. Biol. Soc. Washington, 16: 42. Pulau Lampi 
(= Sulli\'ans Island), Mergui Archipelago. 

Tr.xgulus jav.-^mcus \vili.ia.msom Kloss, 19 1 6 

1916. Tragulus kanchil uilliamsotn Kloss, J. N.H. Soc. Siam, :?.■ 88. Me Song Forest, 
Pre, Northern Siam. 

Tragulus j.wanicus .\ngustiae Kloss, 1918 

1918. Tragulus kanchil angustiae Kloss, J. Fed. Malay States Mus. j: 254. Bankachon, 
\'ictoria Point, Tenasserim. Ranges to extreme north of Lower Siam. 

Tr.agulus javanicus merg.atus Thomas, 1923 

1923. Tragulus ravus mergalus Thomas, J. Bombay N.H. Soc. 2g: 85. King's Island, 
Mergui Archipelago. 


Genera: Alces, page 373 Elaphurus, page 370 

Axis, page 360 Hydropotes, page 354 

Capreolus, page 371 Moschus, page 353 

Cervus, page 361 Muntiacus, page 355 

Dama, page 358 Rangifa, page 375 
Elaphodui, page 357 

Amongst living Ccr\idae Moschus and Hrdrnpotcs stand apart from the remainder 
on account of their lack of antlers. Simpson (1945) divided the living Cerv'idae into 
four subfamilies: the Moschinae, for Moschus alone; the Muntiacinae, for Munliacus 
and Klaphodus ; the Cervinac, for Cenus, Axis, Dama and Elaphurus ; and the Odocoileinae 
in which he recognizes se\eral "tribes" in the Palacarctic, each containing a single 
li\ing genus: Capreolus, Alcc.\, Rangijer and Hydropotes. \lost of this classification is 
foreshadowed in Lydekkcr, and other earlier works. It is here followed, with the. 
exception that wc prefer to follow G. Allen and many others in giving Hydropotes 
subfamily rank. There is a wide evolutionary difi'erencc between primitive deer of 
this description and deer in which antlers are present. 

In the generic di\ision ol the deer we follow Simpson, thereby, perhaps, appearing 
rather conservative to those workers who recognize some five other genera which 
^eem best regarded as subgeneric groups. (See also Pocock, 1923, Classification of the 
C:rr\idae, P.~.S., London, 206.) 


Subfamily Moschinae 

Genus MOSCHUS Linnaeus, 1758 

1758. Moschus Linnaeus, Syst. Nat. loth ed. /; 66. Moschus moschiferus Linnaeus. 

(Opinion 75, International Commission on Zoological Nomenclature.) 
1848. Odontodorcus Gistel, Naturgesch. Thierreichs, 82. Moschus moschiferus Linnaeus. 

I species: Moschus moschiferus, page 353 

Moschus moschiferus Linnaeus, 1 758 Musk Deer (Kastura) 

Approximate distribution of species: in the U.S.S.R., the Altai, the whole of the 
mountain-taiga part of Siberia from the Yenesei up to and including the eastern 
slope of the Kolyma Range (but not occurring in North-Eastern Siberia nor 
Kamtchatka), the Sea of Okhotsk and Sakhalin, Ussuri region. Mongolia, Man- 
churia, Korea, Tibet; in China, Szechuan, Shensi and Shansi, Kansu and possibly 
(? or formerly) Chihli ; Kashmir eastwards to Nepal and Sikkim (Assam and Northern 
Burma, Tate (1947).) (Earlier authors, e.g. Trouessart, quoted the species from 
Indo-China, but this appears doubtful; possibly the result of confusion with a 
Traguloid ?) 

Moschus moschiferus moschiferus Linnaeus, 1758 

1758. Moschus moschiferus Linnaeus, Syst. Nat. loth ed. /.• 66. "Tartary, approaching 

1830. Moschus altaicus Eschscholtz, Isis (Oken), 606. Mongolia. 
1839. Moschus chrysogaster Hodgson, J. Asiat. Soc. Bengal, 8: 203. Nepal. 
1839. Moschus leucogaster Hodgson, loc. cit. Nepal. 
1839. Moschus saturatus Hodgson, loc. cit. Nepal. 

1872. Moschus moschiferus maculatus Gray, Cat. Rum. Mamm. B.NL 96. 
1872. Moschus moschiferus fascia^us Gray, loc. cit. 
1 872. Moschus moschiferus concolor Gray, loc. cit. These names were based on vernacular 

names of Milne-Edwards, 1864, Ann. Sci. Nat. Zool. 2: 62. 
1915. Moschus cacharensis Lydekker (ex Hodgson MS.), Cat. Ung. Mamm. B.M. 4: 

6. Kachar inom. nud.). 
Range: Altai and Sayan Mountains, Siberia and Mongolia. According to Lydekker, 
the Indian Himalayan form is the same and he did not retain the next, which 
Bobrinskii says is of doubtful validity: 

Moschus moschiferus sibiricus Pallas, 1779 

1779. Moschus sibiricus Pallas, Spic. Zool. /j.- 29. Stanovoi Range, Transbaikalia. 

Moschus moschiferus sifanicus Btichner, 1891 

1 89 1. Moschus sifanicus BUchner, Melanges Biol. St. Petersb. 75.- 162. Southern 

Kansu, China. 
1929. Moschus berezovskii Flerov, C.R. Acad. Sci. U.R.S.S. ig28A: 519. Ho-tsi-how 

Pass, near Lungan, Szechuan, China. (Status /rff G. Allen.) 
Range: Kansu, Shensi, Szechuan, in China. 



19! I. Moschus parvipes Hoilister, Proc. Biol. Soc. Washington, 24: i. Mountains near 
Mok-po, South Tscholla Province, Korea. Ranges to Manchuria, and the 
Amur-Ussuri region of Eastern Siberia. 

Moschus moschiferus arcticus Flerov, 1929 

1929. Moschus moschiferus arcticus Flerov, C.R. Acad. Sci. U.R.S.S. ig28A: 516. Mt. 
Toulaiakh-khaia, North-Eastern Taskhaiakhtakh Range, Verhoiansk dis- 
trict, North-Eastern Siberia. 

Moschus moschiferus sachalinensis Flerov, 1929 

1929. Moschus moschiferus sachalinensis Flerov, C.R. Acad. Sci. U.R.S.S. igsSA: 517. 
Sakhalin Island, Eastern Siberia. 

Moschus moschiferus turowi Zalkin, 1945 

1945. Moschus moschiferus turowi Zalkin, C.R. Acad. Sci. U.R.S.S. 46: 331-332. 
Sikhote-Alin National Park, Terney Bay, Amurland. 

SuBFA.MiLY Hydropotinae 

Genus HYDROPOTES Svvinhoe, 1870 

1870. Hydropoles Svvinhoe, P.Z.S. 90. Hydropotes inerrnis Svvinhoe. 
1898. Hydrelaphus Lydekker, Deer of all Lands, 219. Substitute for Hydropoles, 
thought to be preoccupied by Hydropota Rondani, 1861. 

I species: Hidropotes inerrnis, page 354 

Hydropotes inerrnis Svvinhoe, 1870 Chinese Water-Deer 

Appro.ximate distributicn of species: China, the eastern Yangtze Basin, westwards 
to Hupeh. Korea. 

Hydropotes inermis inermis Swinhoe, 1870 

1870. Hydropotes inermis Swinhoe, P.Z.S. 89. Deer Island, in the Yangtze River, a 

few miles upstream from Chinkiang, Kiangsu, China. 
1872. Hydropotes affinis Brooke, P.Z.S. 524. Yangtze River, about 40 miles from 

Shanghai, China. 
1905. Hydropotes kreyenbergi Hilzheimer, Zool. Anz. 2g: 298. Chinkiang, Kiangsu, 

Range: Eastern Yangtze Basin, China. 

Hydropotes inermis argyropus Heude, 1884 

1884. Hydropotes argyropus Heude, C.R. Acad. Sci. Paris, g8: 1017. Hilzheimer, 1906, 
Abh. Mus. Nat. u. Heimatk., Magdeburg, /.• 171. Korea. (Trouessart, 
1898, Cat. Mamm. z: 865, states, erroneously, that Heude's name was a 

nnmen nudum.) 


Subfamily Muntiacinae 

Genus MUNTIACUS Rafinesque, 1815 

18 1 5. Muntiacus Rafinesque, Analyse de la Nature, 56. Cenms muntjak Zimmermann 

(see page 4). 

1816. Cervulus Blainville, Bull. Soc. Philom. Paris, 74. Cervus muntjak Zimmermann. 
1825. Munijaccus Gray, Ann. Phil. 10: 342 (nom. nud.). 

1827. Stylocerus H. Smith, Griffith's Cuvier Anim. Kingd. j.' 319. Cervus muntjak 

1837. Prox Ogilby, P.Z.S. 1826: 135. Prox moschatus Ogilby = Cervus muntjak 

1843. Muntjacus Gray, List. Spec. Mamm. B.M. 173. Cervus muntjak Zimmermann. 
1923. Procops Pocock, P.Z.S. 207. Cervulus feae Thomas & Doria. 

5 species: Muntiacus crinifrons, page 357 
Muntiacus feae, page 357 
Muntiacus muntjak, page 355 
Muntiacus reevesi, page 356 
Muntiacus rooseveltorum, page 356 

Pocock separated M.feae generically on account of the absence of frontal glands. 
Lydekker stated that these were also absent in M. crinifrons, but G. Allen, 1940, 
Mammals of China and Mongolia, 2: 1160, says that they are present in this species, 
and Thomas and Doria say that feae is closely related to crinifrons. Neither is well 
known. Osgood (1932) reviewed the genus and recognized the long-standing species 
listed here, and gave certain colour details and cranial characters to separate AI. 
reevesi from A/, muntjak. He also described a new species, M. rooseveltorum, based on a 
single specimen, which from description appears valid; it seems curiously inter- 
mediate between muntjak and reevesi, being intermediate in size, having the colour 
more as reevesi and the relatively small preorbital pit of muntjak. But the possibility 
that rooseveltorum is a hybrid between muntjak and reevesi is perhaps unlikely, as reevesi 
is unknown from Indo-China, though many mammals from the habitat of 
reevesi, Southern China, do extend into Indo-China. Besides this, rooseveltorum is des- 
cribed as having highly-developed glandular brushes on either side of the chin, 
which Osgood says are usually present in the other species, though much less well 

Muntiacus muntjak Zimmermann, 1780 Indian Muntjac (Barking Deer) 

Approximate distribution of species: Yunnan and Hainan, in Southern China; 
Burma, Assam, Nepal, Peninsular India, Ceylon. Indo-China, Siam, Malay States, 
Sumatra, Java, Borneo, and some adjacent small islands. 

(Muntiacus muntjak muntjak Zimmermann, 1780. Extralimital) 
1780. Cervus muntjak Zimmermann, Geogr. Gesch. 2: 131. Java. 




1785. Cervus vaginalis Boddaert, Elench. Anini. /; 136. Bengal. 

1833. Cerim rativa Hodgson, Asiatick Res. 18, 2: 139. Nepal. 

1840. Cervus mdas Ogilby, in Royle, Illustr. Bot. Himalaya, Ixxiii. India; a mclanistic 

1845. Cervus stvloct-ros Schinz, Synop. Mamm. l': 54(). Renaming of mc/rti. 
1852. Stylocerus munljacus Kelaart, Prod. Faun. Zeylan, 85. Renaming of r(7i;;«fl/». 
Range: Knniann In Bhutan Duars and Chindvvin, Burma: Yunnan, Northern Indo- 

McN'TiACUs MUNTj.xK Ai'REVs H. Smith, 1826 

1826. Cervus aureus H. Smith, Griffith's Cuvier Anim. Kingd. ^: pi. opposite p. 148 

(text, 148, 1827). "Some part of Southern India" (Lydekker, 1915). 
1844. Cervus albipes Wagner, Schreb. Saugeth. Suppl. 4: 394. Bombay and Poona. 
1872. Cervulus tamuUcus Gray, Cat. Ruminants B.M. 94. Deccan, India. 
Range: southern part of Peninsular India. 


1872. Cervulus eurvoslvlis Gray, Cat. Ruminants B.M. 94. Pachebon, Siam. 

MuNTiACUS MUNTJAK GRANDicoRNis Lydekker, 1904 

1904. Cervulus muntjac grandieornis Lydekker, Field, 104: 780. Thouagyen Forest, 
Amherst district, Burma. Range: Burma and Tenasserim. 

MuNTiACus MUNTJAK MALABARicus Lydekker, 191 5 

191 5. Muntiaeus muntjak malabaricus Lydekker, Cat. Ungulate Mamm. B..M. 4: 24. 
Nagarhol, Coorg, Southern India. Range: Malabar coast and Ceylon. 


1928. Muntiaeus muntjak annamensis Kloss, Ann. Mag. N.H. /.• 399. Langbian Peak, 
Southern ,\nnam, Indo-China. 


1930. Muntiaeus muntjak nigripes G. Allen, Amer. Mus. Nov. 430, 11. Nodoa, Island 
of Hainan. Range includes Annam (part). 

Muntiaeus rooseveltorum Osgood, 1932 

Distribution: only known from the type locality, in Indo-China. 


1932. Muntiaeus rooseveltorum Osgood, Field Mus. Publ. Zool. 18: 332. Muong Yo, 
Laos, Indo-China. 

Muntiaeus reevesi Ogilby, 1839 Reeves' Muntjac 

.\ppnjxiniatc distribution of species: Szechuaii, Hupch, eastwards to Fukien and 
adjat ent states in Southern China; ? Formosa. 


1839. Cenms reevesi Ogilby, P.Z.S. 1838: 105. Near Canton, Kwantung, Southern 

1872. Cervulus lachrymans Milne-Edwards, Nouv. Arch. Mus. H.N. Paris, 7, Bull.: 93. 

Moupin, Szechuan, China. 

1873. Cervulus iclateri Swinhoe, P.Z.S. 814. Near Ningpo, Chekiang, Southern China. 

1905. Cervulus sinensis Hilzheiiner, Zool. Anz. sg: 297. Probably Hwei Shan, Anhvvei, 

Southern China. 

1906. Cervulus reevesi pingshiangicus Hilzheimer, Abh. Mus. Nat. u. Heimatk. Magde- 

burg, /.• 169. Pingshiang, Anhwei, China. 
1910. Cervulus bridgemani Lydekker, Abstr. P.Z.S. 38; 191 1, P.Z.S. igio: 989. Hwei 

Shan, Anhwei, China. 
1915. Munttacus lachrymans teesdalei Lydekker, Cat. Ungulate Mamm. B.M. 4: 27. 

Tatung, Yangtze Valley, China. 
Range: as in the species, except Formosa. 

MuNTiAcus REEVESI MicRURUS Sclater, 1875 

1875. Cervulus micrurus Sclater, P.Z.S. 421, pi. 51. ? Formosa. Perhaps a synonym of 
the typical race. 

We follow G. Allen in referring all named forms to the synonymy of the typical 
race, except the last. Lydekker divided this group into three distinct species, and 
several races. 

Muntiacus crinifrons Sclater, 1885 Black Muntjac 

Approximate distribution of species: known from three specimens only, from the 
State of Chekiang, in South-Eastern China. - 

Muntiacus crinifrons Sclater, 1885 

1885. Cervulus crinifrons Sclater, P.Z.S. i, pi. i. Near Ningpo, Chekiang, South- 
Eastern China. 

Muntiacus feae Thomas & Doria, 1889 Fea's Muntjac 

Approximate distribution of species: known only by very few specimens from 
Tenasserim and Siam. 

Muntiacus feae Thomas & Doria, 1889 

1889. Cervulus feae Thomas & Doria, Ann. Mus. Stor. Nat. Genova, y: 92. Thagata 
Juva, south-east of Mt. Mulaiyit, Tenasserim. 

Genus ELAPHODUS Milne-Edwards, 1872 

1872. Elaphodus Milne-Edwards, Nouv. Arch. Mus. H.N. Paris, 7, Bull. : 93. Elaphodus 
cephalophus Milne-Edwards. 

1874. Lophotragus Swinhoe, P.Z.S. 453. Lophotragus michianus Swinhoe. 

I species: Elaphodus cephalophus, page 358 

This genus is closely allied to Muntiacus; its characters are given in Lydekker, 
1915, Cat. Ungulate Mamm. B.M. 4: 34. 



Elaphodus cephalophus Milne-Edwards, 1872 Tufted Deer 

Approximate distribution of species: Szechuan, Hupch, Yunnan, Fukien and 
Cfiekiang in Southern China; Northern Burma. 

Elaphodus cephalophus cephalophus Milne- Edwards, 1872 

1872. Elaphodus cephalophus Milne-Edwards, Nouv. Arch. Mus. H.N. Paris, 7, Bull.: 
93. Moupin, Szechuan, China. Range: to Yunnan and Northern Burma. 

Elaphodus cephalophus michianus Swinhoe, 1874 

1874. Lophotragus michianus Swinhoe, P.Z.S. 453, pi. 59. Near Ningpo, CUiekiang, 

Southern China. 
1904. Elaphodus michianus fociensis Lydekker, P.Z.S. igo^, 2: 169. Fing-ling, Fokien, 

Southern China. 
Range: South-Eastern China. 

Elaphodus cephalophus ichangensis Lydekker, 1904 

1904. Elaphodus ichangensis Lydekker, P.Z.S. Jgo^, 2: 169. Ichang, Hupeh, China. 
Ranges into Szechuan. 

Subfamily C e 

r V 1 n a e 

Lydekker (191 5) gives a key to the genera. He regarded Axis as a subgenus of 
Cervus, to which he gives a key of subgenera on p. 48. Pocock and others, including 
Simpson, have raised Axis to generic rank. 

Genus DAMA Frisch, 1775 

1775. Dama Frisch, Natur-syst. der Vierfiiss. Thiere, 3. Cervus dama Linnaeus. (See 

past- 3.) 
1780. Platvceros Zimmermann, Geogr. Geschichte, 2: 128. Platyceros plinii Zimmer- 

mann = Cervus dama Linnaeus. 
1827 Dama H. Smith, Griffith's Cuvier Anim. Kingd. Mamm. j.- 306. Cervus dama 

1844. Platyceros Wagner, Schreb. Saugeth. Suppl. 4: 340. Cervus dama Linnaeus. 
1855. Dactyloceros Wagner, loc. cit. 5.- 349, 352. Substitute for Dama and Platyceros. 
1893. Machlis Zittcl, Handb. Paleont. 4: 402. (Synonym oi Dama in part, teste Kaup.) 
1898. Palmalus Lydekker, Deer of all Lands, 125. Substitute for Dama. (For use of 

the name Dama, see J. Mammal, jjo, 1949: 94.) 

2 species: Dama dama, page 359 

Dama mesopotamica, page 359 

Dama mesopotamica is provisionally accorded specific rank on account of its greater 
size, the peculiarity of the antlers, and, more especially, on account of the shape of 
the nasals which are much broader across the proximal end than in dama (Brooke, 
1876, gives this measurement as 74 mm. in mesopotamica against 46 mm. in dama). At 



the same time it should be borne in mind that mesopotamica has been found in large 
numbers in the Pleistocene of Palestine, where its antler shape shows great variation. 
The recent range of dama extended to Palestine, and it may well have been that 
within recent times the range of mesopotamica abutted on that of dama and that the 
former should be regarded as a geographical race of the latter. 

Dama dama Linnaeus, 1 758 Fallow Deer 

Approximate distribution of species: the original home is said to be the Mediter- 
ranean region of Southern Europe and Asia Minor, but fallow deer have been 
widely introduced and are now to be found wild in most parts of Western Europe, 
the Western Ukraine and Baltic States. Introductions were made in North Africa, 
but it is doubtful whether there are any there established wild. The present status in 
Asia Minor is obscure. 

Dama dama Linnaeus, 1 758 

1758. Cervus dama Linnaeus, Syst. Nat. loth ed. /.• 67. Sweden (introduced). 

1780. Platyceros plinii Zimmermann, Geogr. Gesch. 2: 128. Renaming oi dama. 

1798. Cervus platyceros Cuvier, Tabl. Elem. H.N. Anim. 160. Renaming o^ dama. 

1816. Cervus mauricus Cuvier, Bull. Soc. Philom. Paris, 72. No locality. (Melanistic.) 

1829. Cervus dama var. vulgaris Fischer, Syn. Mamm. 448. 

1829. Cervus dama var. leucaethiops Fischer, loc. cit. (albino). 

1829. Cervus dama var. maura Fischer, loc. cit. Renaming o{ mauricus. 

1874. Dama platyceros niger Fitzinger, S.B. Akad. Wiss. Wien, 65, i: 553. 

1874. Dama platyceros varius Fitzinger, loc. cit. 555. 

1874. Dama platyceros albus Fitzinger, loc. cit. 555. (These names based on melanistic, 

spotted and albino variations.) 

Occurs in Spain, France, United Kingdom, Holland, Belgium, Denmark, 
Switzerland, Italy, Austria, Germany, Poland, Czechoslovakia, Hungary, the Baltic 
States, Norway, Sweden, the Ukraine, and Island of Rhodes. Probably throughout 
the Balkans as well. 

Dama mesopotamica Brooke, 1875 Persian Fallow Deer 

Approximate distribution of species: Persia, and adjacent parts of Iraq. This 
deer may now be extinct. A male was obtaiiied on 21 July 1917, at Zakho, 37°o8' N., 
42°37' E. (Northern Iraq), and another specimen has been recorded from the 
Juanrud district, north of Kermanshah, Western Persia. The last recorded specimen 
from the Luristan district appears to have been one seen in the upper reaches of the 
River Diz, about 1906. (See also Brooke, P.Z-S., London, i8y6: 298, and 1878: 790, 
Bate, 1937, The Stone Age of Mount Carmel, i, 2: 210, and Pocock, 1946, J. Soc. Pres. 
Fauna Emp. 53: 53.) 

Dama mesopotamica Brooke, 1875 

1875. Cervus [Dama) mesopotamicus Brooke, P.Z.S. 264. Luristan Province of Persia. 
1905. Cervus dama mesopotamiae Trouessart, Cans. Sci. Soc. Zool. France, /.■ 405. 



Genus AXIS H. Smith, 1H27 

1827. Axis H. Smith, Griffith's Cuvier, Ariim. Kingd. 5.- 312. Cervus axis Erxleben. 
1846. Hvelaphus SuikIcnciH, K. Svenska Vctcnsk. Akad. Handl. 1844: 180. Cervus 
porcinus Zimmrinuuin. \'aiid as a subgenus. 

2 species in the area covered by this list: 
Axis axis, page 360 
Axis porcinus, page 360 

The latter is here separated subgenerically as Hvelaphus, and in this we fnilow 
Simpson; Pocock, 1943, J. Bombay JS'.H. Soc. 44: 174, gave it generic rank. 

Subgenus AXIS H. Smith, 1827 

Axis axis F.ixlrbcn, 1777 Clhital, Axis Deer, Spotted Deer 

Approximate distribution of species: Ceylon and Peninsular India, northwards to 
Kumaon, Nepal, Sikkim, Bengal. 

Axis AXIS .AXIS Erxleben, 1777 

1777. Cervus axis Erxleben, Syst. Regn. Anini. 312. Banks of the Ganges, India. 

1792. Cervus axis maculatus Kerr, Anim. Kingd. 300. Banks of the Ganges. 

1829. Cervus axis var. indicus Fischer, Syn. Mamm. 619. 

1 83 1. Cervus nudipalpehra Ogilby, P.Z.S. 1830-31: 136. Banks of the Ganges. 

1842. Axis major Hodgson, J. Asiat. Soc. Bengal, 10: 941. 

1842. Axis minor Hodgson, loc. cil. 

Axis AXIS cEVLONENSis Fischer, 1829 

1829. Cervus axis var. cevlonensis Fischer, Syn. Mamm. big. Ceylon. 

1905. Cervus iRusa) axis zeylanicus Lydekker, Field, io§: 947. 

Subgenus HYELAPUL'S Sunde\all, 1846 

Axis porcinus Zimmermann, 1780 Hog Deer (Para) 

Approximate distribution of species: from Sind and the Punjab, through Kumaon, 
Nepal and Bengal to Assam, Burma, Indo-C^hina and Siam. Not found in Peninsular 
India but in Ceylon, where it is said to have been introduced by the Dutch or 

Axis PORCINUS poRc:iNus Zimmermann, 1780 

1777. Cervus porcinus Zimmermann, Spec. Zool. Geogr. 532. Bengal. (Zimmermann 

[I'j'J'J) is not an available work (Bull. Zool. Nomencl. 1950, 4: 547) ). 
1780. Cervus porcinus Zimmermann, Geogr. Gcsch. 2: 131. Bengal. 
1784. Cervus porcinus Schreber, Saugeth. 5, pi. 251. Bengal : based on a specimen 

belonging to Lord Clive and described by Pennant, 1771). 
(?) 1827. Cervus pumilio H. Smith, Griffith's Catvier Anim. Kingd. 4: 120. Locality 




(?) 1852. Axis oryzus Kelaart, Prodr. Faun. Zeyl. 83. Ceylon. Regarded by Pocock 

(1943) as a valid race. 
1883. Cervus minor Sclater, List Anim. Zool. Gardens, 169; not of Hodgson, 1842. 


Range: Indian range of the species above. 

Axis porcinus annamiticus Heude, 1888 

1888. Hyelaphiis annamiticus Heude, Mem. H.N. Emp. Chin, 2: 50. Baria, Indo-China. 
1908. Cervui porcinus hecki Lydekker, Field, ///.■ 583. Siam. 

Genus CERVUS Linnaeus, 1758 

1758. Cervus Linnaeus, Syst. Nat. loth ed. /.• 66. Cervus elaphus Linnaeus. 

1827. Rusa H. Smith, Griffith's Cuvier Anim. Kingd. 4: 105. Cervus unicolor Kerr. 

Valid as a subgenus. 
1827. Elaphus H. Smith, Griffith's Cuvier Anim. Kingd. j.' 307. Cervus elaphus 

1838. Harana Hodgson, Ann. N.H. /.• 154. Cervus wallichii Cuvier. 
1838. Rucervus Hodgson, Ann. N.H. /.■ 154. Cervus elaphoides Hodgson = Cervus 

duvaucelii Cuvier. Valid as a subgenus. 
1841. Pseudocervus Hodgson, J. Asiat. Soc. Bengal, 10: 914. Cervus wallichii Cuvier. 
1843. Panolia Gray, List. Mamm. B.M. 180. Panolia acuticornis Gray = Cervus eldii 

M'Clclland. Valid as a subgenus. 
1846. Hippelaphus Sundevall, K. Svenska Vetensk. Akad. Handl. 1844: 177. Not of 

Reichenbach, 1835. Cervus hippelaphus Cuvier. 
1846. Strongyloceros Owen, Brit. Foss. Mamm. Birds, 470. Cervus elaphus Linnaeus. 
1870. Sika Sclater, P.Z.S. 115. Cervus sika Temminck = Cervus nippon Temminck. 

Valid as a subgenus. 
1872. Pseudaxis Gray, Cat. Ruminants B.M. 70. Cervus taiouanus Blyth (a race of 

C. nippon Temminck). 
1874. Elaphoceros Fitzinger, S.B. Akad. AViss. Wien, 68, i: 347, 352. Cervus sika 

Temminck = Cervus nippon Temminck. 
1888. Samhur Heude, Mem. H.N. Emp. Chin. 2: 8. Cervus arislotelis Cuvier. 
1898. Sikaillus Heude, Mem. H.N. Emp. Chin. 4: 98. Cervus sika Temminck. 

1898. Sica Trouessart, Cat. Mamm. 878. (Substitute for Sika.) 

1899. Eucervus Acloque, Faune de France, Mamm. 71. Not of Gray, 1866. Cervus 

elaphus Linnaeus. 
1930. Przewalskium Flerov, C.R. Acad. Sci. U.R.S.S. 1 15. Cervus albirostris Przewalski. 

Valid as a subgenus. 
1943. Thaocervus Pocock, J. Bombay N.H. Soc. 43: 554, 559. Rucervus schomhurgki 

Blyth. Valid as a subgenus. 

7 species in the area covered by this list: 

Cervus albirostris, page 366 Cervus nippon, page 364 

Cervus duvauceli, page 363 Cervus schomhurgki, page 363 

Cervus elaphus, page 367 Cervus unicolor, page 362 
Cervus eldi, page 364 



Each of these deer has a subgeneric name. Formerly, eldi, schomburgki and dtivauceli 
were referred to Rucenms, but Pocock, 1943, J. Bombay N.H. Soc. ^j: 553, in reviewing 
the group, separated the three species into three genera. See also Pocock, 1942, 
7. Bombay „V.//. Soc. 42' -9^' fo'" ^ review of Indian Cervus sensii strkto compared with 
Przewahkium which contains the single species alhiroslris. 

Lydekker retained three species in the subgenus Sika and seven in Cervus sensu 
strkto, but we ha\T reduced them to one each. Many of the names which have been 
given to deer are based on antler differences which modern observations have 
shown to be well within the range of phenotypical variation, due to differences of 
feed. The non-genetic nature of much of this variability is well illustrated by the case 
of the British Red Deer which were imported into New Zealand (see Hu.xley, 1931, 
P.^.S. 832). Here they soon came to resemble Carpathian Red Deer, but after a 
time, when the feed deteriorated on account of other activities of man, the deer 
"went back" and in the end came once more to resemble the small-sized deer, with 
relatively poorly developed antlers, which had been their starting point. 

Lydekker gives the characters of the species here retained. 

Subgenus RUSA H. Smith, 1827 

Cervus unicolor Kerr, 1 792 Sambar 

Approximate distribution of species: Szechuan, Yunnan, Kwantung, Hainan, 
Formosa. Ceylon, northwards through Peninsular India to Kumaon and Nepal, 
Assam, Burma. Indo-China, Siam, Malay States, Sumatra, Java, Borneo, Celebes, 
the Philippines and many small Malayan islands. 

Cervus unicolor unicolor Kerr, 1792 

1792. Cervus axis unkolor Kerr, Anim. Kingd. 300. Ceylon (as restricted by Hamilton 

1792. Cervus axis major Kerr, Anim. Kingd. 300. Ceylon. 

1799. Cervus albicornis Bechstein, Uebcrs. vierf Thiere, /.• 112. Substitute for major. 
1898. Cervus unicolor typicus Lydekker, Deer of all Lands, 146. 
Range: Ceylon i Pocock, 1943). 

Cervus unicolor nicer Blainville, 1816 

1 816. Cervus niger Blainville, Bull. Soc. Philom. Paris, 76. "Probably somewhere in 

North India" (Pocock). 
1823. Cervus aristotelis Cuvier, Oss. Foss. ed. 2, 4: 503. Nepal. 
1823. Cervus leschenaulti Cuvier, Oss. Foss. ed. 2, 4: 506. Coromandel, India. ^ 
1827. Cervus hippelaphus H. Smith, Griffith's Cuvier Anim. Kingd. 4: 105. Not of 

Erxleben, 1777. ? Bengal. 
1 83 1. Cervus jarai Hodgson, Gleanings Science, 3: 321. Nepal. 
1 84 1. Cervus Ac/frarm/j Hodgson, J. Asiat. Soc. Bengal, /o, pi. opposite 722, and 10: 

914 (where the spelling is helerocervus). 
1 84 1. Cervus nepalensis Hodgson, loc. cit. Nepal. 
1843. Axis pennantii Gra.y, List Mamm. B.M. 180. India. 

Range : Peninsular India (apart from western desert and semi-desert areas) to Nepal. 
The name is revived by Pocock, 1943, J. Bombay N.H. Soc. 44: 30. 



Cervus unicolor EquiNus Cuvier, 1823 

1823. Cervus equinus Cuvier, Oss. Foss. ed. 2, 4: 45. Sumatra. 

1861. Cervulus cambojensis Gray, P.Z.S. 138. Cambodia, Indo-China. 

1888. Sambur curvicornis and longkornis, and oiitrevanus Heude, Mem. H.N. Emp. Chin. 
2: 42 ; and S. planidens, S. colombertinus, S. combalbertinus, 43 ; and S. lignarius, 
S. lemeanus, 44 ; and 6'. errardianus, S. joubertianus, S. latidens, S. planiceps, 45 ; 
and S. officialis, S. simoninus, S. brachyrhinus, S. verutus, 46. All from Cochin- 
i8g6. Rusa dejeani Pousargues, Bull. Mus. H.N. Paris, 2: 12. Szechuan, China. Status 

fide Pocock (1943). 
Range: Sumatra, Malay States, Indo-China, Siam, Assam, Burma, Yunnan, 
Szechuan, Hainan. 

Cervus unicolor swinhoei Sclater, 1^62 

1862. Cervus swinhoii Sclater, P.Z.S. 152. Formosa. 

Subgenus RUCERVUS Hodgson, 1838 

Cervus duvauceli G. Cuvier, 1823 Swamp Deer; Barasingha 

Approximate distribution of species: India, north of the Ganges from Kumaon to 
Assam, and south of the Ganges principally in the Central Provinces (Pocock). 

Cervus duvauceli duvauceli G. Cuvier, 1823 

1823. Cervus duvaucelii G. Cuvier, Oss. Foss. ed. 2, 4: 505. "Based on sketches of 
antlers sent by Du\'aucel, locality not recorded but no doubt North India" 

1834. Cervus bahrainja Hodgson, P.Z.S. gg. Nepal. 

1835. Cervus elapho'ides Hodgson, J. Asiat. Soc. Bengal, 4: 648. Substitute for bahrainja. 
1837. Cervus smithii Gray, P.Z.S. 45. The drawing in the British Museum on which 

this name is based is of a duvaucelii with aberrant antlers. Northern India. 
1843. Cervus dimorphe Hodgson, J. Asiat. Soc. Bengal, 12: Sgy. Saul Forests of the 

Morung, Nepal. 
1850. Cervus euceros Gray, Knowsley Menagerie, pi. 40 [euryceros in text, p. 61). India. 
1868. Cervus eudadoceros Falconer, Pal. Mem. /.• 587. West bank of Ganges, south of 

Hardwar, United Provinces, India. 
Range: north of the Ganges, India. 

Cervus duvauceli branderi Pocock, 1943 

1943. Rucervus duvaucelii branderi Pocock, J. Bombay N.H. Soc. 4;^: 558. Mandla, 

Central Provinces, India. Range : south of the Ganges, Central Provinces, 


Subgenus THAOCERVUS Pocock, 1943 

Cervus schomburgki Blyth, 1863 Schomburgk's Deer 

Appro.ximate distribution of species: Siam, if not now extinct. 



Cervus schomburgki Blyth, 1863 

1863. Cervus or Rucervus schomburgki Blyth, P.Z.S. 155. Siam. 

Subgenus PAMOIJA Gray, 1843 

Cervus eldi M'C:lclland, 1842 Thamin, or Eld's Deer 

Approximate distribution of species: Manipur, Burma, Hainan, Siam, Indo-China. 
(Thomas, 1918, J. Bombay JV.H. Soc. 2j: 365, says that all references to Formosa in 
relation to British Mu§cum specimens of this deer should be deleted and replaced by 
Hainan; the error is attributed to Gray or Gerrard.) 

Cervus eldi ei.di M'Cielland, 1842 

1842. Cervus eldii M'Clclland, Calcutta J.N. H. i'.- 417. Manipur, Assam. 

1843. Cervus (Rusa) frontalis M'Clelland, Calcutta J.N. H. j.- 401. Renamins; of eldti. 
1843. Panolia acuticornis Gray, List Mamm. B.M. 180. Manipur. 

1845. Cervus lyralus Schinz, Synop. Mamm. 2: 395. Based on M'Clclland (1841, 
which was a description without name). 

1864. Panolia acuticauda Blyth, P.Z.S. i86j: 370. Renaming otjrontalls. 
1898. Cervus eldi typicus Lydekker, Deer of all Lands, 200. Manipur. 
1901. Cervus eldi cornipes Lydekker, Nature, §^: 257. Manipur. 

Range: Manipur. 

Cervus eldi siamensis Lydekker, 19 15 

1915. Cervus eldi siamensis Lydekker, Cat. Ungulate Mamm. B.^L 4: 104. Southern 

Siam. Renaming oi platyceros Gray, 1843. 
1843. Panolia platveeros Gray, List Mamm. B.NL 181. Siam. Not Cervus platrceros 

Cuvier, 1 798. 
(?) 1918. Rucervus plalveeros hainanus Thomas, J. Bombay X.H. Soc. 2^: 364. Hainan 

Island, Southern China. 

Range: Indo-China, Siam, Hainan. 

Cervus eldi th.amin Thomas, 1918 

1918. Rucervus thamin Thomas, J. Bombay N.H. Soc. 2fj: 364. Pegu, Burma. 

1918. Rucervus thamin briicei Thomas, loc. cit.: 366. Thimbaung-Gwin Plain, Ruby 

Mines district. Upper Burma 
Range: Burma, Tenasserim, Siam (in part). 

Subgenus ,S7AV1 Si later, 1870 

Cervus nippon IVmminck, 1838 Sika Deer, Japanese Deer 

Appro.ximate distribution of species: .Southern LJssuri district of Eastern Siberia; 
Japan, Manchuria, Formosa; in China, Chihli, Shansi, and the eastern Yangtze 
Basin from CJhekiang and Kiangsu into Northern Kwantung. 



Cervus NIPPON NIPPON Tcmniinck, 1838 

1838. Cervus nippon Temminck, Coup d'oeil sur la faune des iles de la Sonde et de 
rempire dujapon, xxii. Japan. 

1845. Cervus sika Temminck, Fauna Japonica, Mamm. 54, pi. 17. Japan. 

1846. Cervus {Hippelaphus) japonicus Sundevall, K. Svenska Vetensk. Akad. Handl. 

1844: 178. Japan. 

1878. Cervus mantchuricus minor Brooke, P.Z.S. 909. Japan. Nee Wagneir, 1855. 

1884. Sika schlegeli Heude, Cat. Cerfs Tachetes, y-jfuscus, 7, hollandianus, 8, all from 
"the small islands south of Japan"; infelix, 7, brachjpus, 8, both from Goto 
Islands, Japan ; orthopus, 8, Kobe, Japan ; blakistoninus , dolichorhinus, legrandi- 
anus, 9, vesoensis, 10, sylvanus, 11, all from Nippon and Yezo; aplodontus, 10, 
north of Tokyo, mitratus, 10, Tokyo; xendaiensis, 11. Sendai, Nippon. 

1888. Sika paschalis Heude, Mem. H.N. Emp. Chin. 2, pi. 18, fig. i; aceros, fig. 2; 

rex, fig. 5; dejardinus, fig. 6; marmandianus, pi. 19, fig. 6; all from Goto Islands, 

1893. Cervus sica Lydekker, Horns & Hoofs, 284. Emendation of sika. 
1897. Cervus sica typicus Lydekker, P.Z.S. 39. 

1897. Sika sendaiensis Heude, Mem. H.N. Emp. Chin, j.- 98 (for xendaiensis 1884), 

schizodonticus, loi, Tokyo; orthopodicus (for orthopus 1884); ellipticus, elegans, 
Sendai; minoensis, 104, Mino, west of Tokyo; rutilus, 105, Yezo (= Hok- 
kaido), Japan. 

1898. Sikaillus daimius Heude, Mem. H.N. Emp. Chin. 4: loi; regulus, 103; sicarius, 

105; consobrinus, 107; lalidens, 108; Goto Islands, Japan. 
Range: Japan (Hondo, Hokkaido, Shikoku, Kiushu, Tsushima, Yakushima) and 

Cervus nippon taiouanus Blyth, i860 

860. Cervus taiouanus Blyth, J. Asiat. Soc. Bengal, sg: 90. Formosa. 

862. Cervus taevanus Sclater, P.Z.S. 152 (for taiouanus Blyth). 

872. Pseudaxis taivanus Gray, Cat. Ruminants, B.M. 70. 

882. Cervus tai-oranus Heude, Bull. Soc. Philom. 6: 184 (emendation). 

882. Cervus devilleanus Heude, loc. cit. 187. Formosa. 

884. Sika dominicanus Heude, Cat. Cerfs Tachetes, 6, novioninus, schulziamis, morrisi- 
anus, 6, all from Formosa. 
Range: Formosa. 

Cervus nippon hortulorum Swinhoe, 1864 

1 86 1. Cervus pseudaxis (?) Gray, P.Z.S. 236. Nee Gervais, 1841. 

1864. Cervus hortulorum Swinhoe, P.Z.S. 169. "Gardens of the Summer Palace, 
Pekin." According to G. Allen, its true home was Manchuria. 

1864. Cervus mantchuricus Swinhoe, P.Z.S. 169. Ying-tzu-kou, Newchwang, Manchuria. 

1874. Cervus euopis Sclater, P.Z.S. 151. Newchwang, Manchuria. 

1876. Cervus dybowskii Taczanowski, P.Z.S. 123. Southern Ussuri district of Man- 

1884. Sika mierospilus Heude, Cat. Cerfs Tachetes, 11. Manchuria. 

1889. Cervus mantschuricus major Noack, Humboldt, 8: 9. Not of Kerr, 1792. 

1894. Sika imperialis Heude, Mem. H.N. Emp. Chin. 2: 146. Manchuria, 
igio. Cervus hortulorum typicus \Vard, Rec. Big Game, ed. 6, 52. Manchuria. 
Range: Korea, Manchuria, and adjacent parts of Eastern Siberia; Quelpart Is. 



Cervus NIPPON MANDARiNus Milnc-Edwards, 1871 

1871. Cnvus mandarinus Milnc-Edwards, Rech. Mamm. 184. Said to have come from 
Northern China. This name is used by G. Allen for the race in Chihli, 
which he remarks is probably "now nearly exterminated". 

1882. Cervus cvdorhinus Heude, Bull. .Soc. Philom. 6: 188; hycmalis, 188; both from 
Shantung Pro\ince, China. 

Cervus nippon kopschi Swinhoe, 1873 

1873. Cervus kopschi Swinhoe, P.Z.S. 574. Kien-chang, Kiangsi, Southern China. 

1882. Cervus frinianus Heude, Bull. Soc. Philom. 6: 185; gracilis, 185; lachryinosus, 

igrwlus, 1 86;- all from right bank of Yangtze, below Lake Poyang; andreanus, 

186; joretianus, 187; both from Ningkwofu, 75 miles south of Nanking, 

Southern China. 
1884. Sika brachvrhinus Heude, Cat. Cerfs Tachetes, 2; cycloceros, 2; gnlloanus, 

pouvrelianus, microdontus, 3; oxycephahis, 4; yuanus, 5; all from right bank of 

Yangtze, below Lake Poyang. 
1888. Sika granulosus Heude, Mem. H.N. Emp. Chin. 2, pi. o, fig. 2; surdesceris, pi. lA, 

fig. 9; no locality. 
1894. Sika riverianus Heude, Mem. H.N. Emp. Chin. 2: 153. Poyang Lake; dugcn- 

neanus, 156; arietinus, 162; no locality. 
Range : eastern parts of Southern China. 

Cervus .nippon grassianus Heude, 1884 

1884. Sika grassianus Heude, Cat. Cerfs Tachetes, 12. Tsinglo-hsien, Northern Shansi, 
China. (Now nearly exterminated, G. Allen.) 

Cervus nippon keram.'^e Kuroda, 1924 

1924. Sika nippon keramae Kuroda, on New Mamm. Riukiu Islands (Tokyo), 12. 
Zamamishifna, Kerama Group, Middle Riukiu Islands. 

Incertae scdis 

1888. Sika munitus Heude, Mem. H.N. Emp. Chin. .', pi. 18, fig. 3; kt'matoceros, pi. 19, 
fig. I ; modestus, pi. 19, fig. ^\fuscus, pi. 19, fig. 5. No localities. 

1924. Cervus matsumolei Kishida, Monogr. Jap. Mamm. 36. {N.V.) Hokkaido. 
"Doubtful form," according to Kuroda. 

Subgenus PR~E]\'ALSKIUM Flerov, 1930 

Cervus albirostris Przcwalski, 1883 Thorold's Deer 

.A.pproximatc distribution of species: Szechuan, Kansu, Tibet and Kuku-nor. 

Cervi;s albirostri.s Przewalski, 1883 

1883. Cervus albirostris Przewalski, Third Journey in C. Asia, 124. Three km. above 

mouth of Kokusu River, western Humboldt Mountains, Nan-Shan, Western 
Kansu, China. 
1883. Cervus sellatus Przewalski, Third Journey in C. Asia, 125. Same locality. 



1889. Cervus dybowskii Sclater, J. Asiat. Soc. Bengal, §8, 2: 186. Not of Taczanowski, 

1876. Bought in bazaar at Darjeeling. 
1893. Cervus thoroldi Blanford, P.Z.S. 444, pi. 34. Two hundred miles north-east of 

Lhasa, Tibet. 

Subgenus CERVUS Linnaeus, 1758 

All named forms are here referred to one species elaphus, as explained in the note 
under the genus Cervus above. 

Cervus elaphus Linnaeus, 1758 Red Deer 

(Wapiti, Hangul, Shou and others included) 

Approximate distribution of species : the Palaearctic region, eastwards to Man- 
churia and Eastern Siberia, south to the Yangtze, and into the Indian region along 
the southern slopes of the Himalayas; Algeria and Tunis, where rare. North 

(In more detail: British Isles and Europe (in parts re-established by man after 
earlier extinction), Portugal, Spain, France, Switzerland, Netherlands to Denmark, 
Norway, Sweden, Poland and Latvia; Germany, southwards to Rumania, Bulgaria 
and Greece; including Corsica, Sardinia and Italy (introduced); Western VVhite 
Russia, Western Ukraine, Crimea, Caucasus, Russian Turkestan, and Southern 
Siberia from Tarbagatai and Altai Mountains to Tartarsk Straits and Sea of Japan, 
northwards roughly to the parallel of the northern tip of Lake Baikal (Bobrinskii) ; 
Asia Minor, Persia, Afghanistan; Zungaria, Mongolia, Manchuria; in China from 
the states of Kansu, Shansi, Szechuan; Tibet; Kashmir, Sikkim and Bhutan. Algeria, 

Cervus elaphus elaphus Linnaeus, 1758 

1758. Cervus elaphus Linnaeus, Syst. Nat. loth ed. /.• 67. Southern Sweden. 
1898. Cervus elaphus typicus Lydekker, Deer of all Lands, 65. 

Range: Sweden. 

Cervus elaphus hippelaphus Erxleben, 1777 

1777. Cervus elaphus hippelaphus Erxleben, Syst. Regn. Anim. /, Mammalia: 304. The 
Ardennes. (Schwarz, 1938, Z. f Sauget. 8: 276.) 

1822. Cervus elaphus germanicus Desmarest, Mamm. 434. The Ardennes. 

1822. Cervus elaphus albus Desmarest, Mamm. 435. Albino form. Nee Kerr, 1792. 

1845. Cervus elaphus albifrons Reichenbach, Vollstand. Naturgesch. Saug. j.- pi. 3 bis, 
fig. 26. (Tame variety.) 

1874. Cervus elaphus varius Fitzinger, S.B. Akad. Wiss. Wien, 6g, i: 574. Germany 
(partial albino). 

1903. Cervus vulgaris Botezat, Morph. Jb. 32: 115. Renaming of elaphus. 

(?) 1903. Cervus vulgaris campestris Botezat, Morph. Jb. 52.- 154. Caipathian Moun- 
tains, Bukowina, Rumania. 

(?) 1903. Cervus vulgaris montanus Botezat, loc. cit. 155. Carpathian Mountains. 



Cervus elaphus hippelaphus [conld.] 

1907. Cervus balticus Matschie, Weidwerk in Wdrt und Bild, 16: 18G. Near Licbe- 

miihl, East Prussia, Germany. 
1907. Cervus alhicus Matschie, loc. cit. Muskau, Oborkiusitz, Silesia, Germany. 
1907. Cervus rhenanus Matschie, loe. cit. Viernhcim, Hessen-Darmstadt, Germany. 
1907. Cervus bajovaricus Matschie, loc. cit. Rohncr, Konigssee, Upper Bavaria, 

1 91 2. Cervus elaphus neglectus Matschie, Deutsche Jager-Zeit,5'?.- 688. Posen, Germany. 
1912. Cervus elaphus visurgensis Matschie, loc. cit. 734. Rhineland, Germany. 
1912. Cervus elaphus debilis Matschie, loc. cit. 734. Rhineland. 
1912. Cervus elaphus saxorucus Matschie, loc. cit. lyj. Saxony, Germany. 
Range: France, Holland, Belgium, Denmark, Itah', Gentral Europe and the Balkans, 

Western Russia. 

Cervus elaphi's corsioanus Erxleben, 1777 

1777. Cervus elaphus var. corsicanus Erxleben, Syst. Regn. Anim. 304. Corsica. 

1822. Cervus mediterraneus Blainville, J. Physique, ^4: 262. Corsica. 
1848. Cervus corsiiiiacui Gervais, Ann. Sci. Nat. Zool. 10: 206. Corsica. 

1855. Cervus elaphus niinar Wagner, Schreb. Siiugeth. Suppl. 5.' 354. Substitute for 

corsicanus . 
Range; Corsica, Sardinia. 

Cervus elaphus wallichi Cuvier, 1823 Shou 

1823. Cervus wallichii G. Cuvier, Oss. Foss. ed. 2, 4: 505. Nepal (or probably 

Mansarowar Lake, Nari-Khorsum district, Tibet, according to Lydekker). 
1841. Cervus qffinis Hodgson, J. Asiat. Soc. Bengal, 10: 721, Saul Forest, Nepal. 

1850. Cervus tibetanus Flodgson, J. Asiat. Soc. Bengal, ig: 466. Lingmo, Phari, 

Dingcham, Tibet. 

1 85 1. Cervus narivanus Hodgson, J. Asiat. Soc. Bengal, 20, pi. 8. Western Tibet. 
Range: Chumbi \'alley (Southern Tibet), Bhutan and Tibet; under the name affuiis 

listed by Bobrinskii from Russian Middle Asia, Northern Afghanistan, the Amu- 

Cervus el.-xphus barbarus Bennett, 1848 Barbary Stag 

1 048. Cervus barharus Bennett, List .'\nim. Gardens Zool. Soc. London, 1833: 48 
(nom. mid.); 1848 (February), Bennett in F'raser, Zool. Typica, pi. 13. 
Ttmis. Range: the Barbary Stag is now tonfmed to a strip ol' forest 
country on the Algcrian-Timisian border. 

C^ERVus elaphus hanglu Wagner, 1844. Hangul; Kashmir ''Barasingha" 

1844. Cervus hanglu \Vagner, Schreb. Saugeth. Suppl. 4: 352 (footnote). Kashmir. 

1847. Cervus casperianus Gray, List Osteol. Specimens B.M. 747. Kashmir. Gray 
wrote "cashmerensis" and then, intending to emend the spelling, he for some 
reason put "casperianus" in the list of errata; this is clearly a lapsus calami. 

1859. Cervus cashmeriensis Adams, P.Z.S. i8§8: 529. 

r868. Cervus cashmeerianus Falconer, Palacont. Mem. /; 576. Kashmir. 

1874. Cervus cashmirianus Fitzinger, S.B. Akad. Wiss. W'ien, 6g, i : 586. 

Range: Kashmir. 



Cervus elaphus maral Gray, 1850. Maral 

1840. Cervus maral Ogilby, Rep. Council Zool. Soc. 22, nom. nud. 

1850. Cervus maral Gray, Knowsley Menagerie, pis. 38, 39. Persia. (These plates are 

of the specimens referred to by Ogilby.) 
1886. Cervus caspius Radde, Fauna u. Flora siidwestl. Caspi-Gebietes, 10. Talysh 

district of Azerbaijan, Transcaucasia. 

1914. Cervus caucasicus Winans, Amer. Mus. J. 14: 67, nom. mid. 

Range: Lydekker regards this deer as ranging west to the Hungarian Carpathians, 
and thought the name montanus of Botezat, 1903, might be the same (see above, 
under synonymy of C. e. hippelaphus) . Crimea, Asia Minor, Northern Persia, 

Cervus elaphus xanthopygus Milne-Edwards, 1867 Manchurian Wapiti 

1867. Cervus xanthopygus Milne-Edwards, Ann. Sci. Nat. Zool. 8: 376. Near Pekin, 

Chihli, C'hina. 
1880. Cervus luhdorjii Bolau, Abh. Naturw. Hamburg, j: 33. Bureatish Steppe, 

Northern Manchuria. 
i88g. Cervus isubra Noack, Humboldt, 8: 12, fig. 5. Based on same specimen as 

1892. Elaphus ussuricus Heude, Mem. H.N. Emp. Chin. 2: 113. Ussuri River. 

1897. Cervus bedfordianus Lydekker, P.Z.S. i8g6: 932. Manchuria. 

1898. Cervus xanthopygus typicus Pousargues, Mem. Soc. Zool. France, //.■ 209. 
Range: Manchuria, Mongolia, Amur-Ussuri region of Siberia. 

Cervus elaphus songaricus Severtzov, 1873 

1873. Cervus maral var. songarica Severtzov, Mem. Soc. Amis. Sci. Nat. Moscou, 
8, 2: 109. Zungarian Tian-Shan, probably near Kuldja, Chinese 

1876. Cervus eustephanus Blanford, P.Z.S. i8j§: 637. Tian-Shan Mountains. 

Cervus elaphus yarkandensis Blanford, 1892 

1 892. Cervus cashmirianus yarkandensis Blanford, P.Z.S. 1 1 7. Maralbashi Forest, Chinese 

Cervus elaphus asiaticus Lydekker, 1898 

1898. Cervus canadensis asiaticus Lydekker, Deer of all Lands, 104. "The district to the 
southward of Lake Teletsk, near the sources of the Yenesei" (G. M. Allen, 

. '93°)- 
1873. Cervus maral var. sibirica Severtzov, Mem. Soc. Amis. Sci. Nat. Moscou, 8. 2: 

109. Siberia. Not Cervus sibiricus Schreber, 1784. 
1907. Cervus biedermanni Matschie, S.B. Ges. Naturf. Fr. 223. Teletskoye Lake, at 
source of Ob, Siberia. 

1915. Cervus canadensis baicalensis Lydekker, Cat. Ungulate Mamm. B.M. 4: 134. 

Sayan and Baikal Mountains, west of Lake Baikal. To replace sibirica 
Severtzov, preoccupied. 
Range: from the Altai to Transbaikalia. 



Cervus elaphus bactrianus Lydekker, 1900 

1900. Cervus bactrianus Lydekker, Ann. Mag. N.H. 5.' 196. Tashkent, Russian 

1904. Cervus hagenbeckii Shitkov, Zool. Jb. Abt. Syst. 20, fig. 4 and p. 103. Russian 


Cervus elaphus wachei Noack, 1902 

1902. Cervus wachei Noack, Zool. Anz. 25.' 146. Shingielt Valley, in neighbourhood 
of the Black Irtish, Kobdo, Western Mongolia. ? Synonym of asiatieus. 

Cervus elaphus atlanticus Lonnberg, 1906 

1906. Cervus elaphus atlanticus Lonnberg, Arkiv. Zool. j, 9: g. Hitteren Island, Trond- 
hjem, Norway. Range: west coast of Norway. 

Cervus elaphus scoticus Lonnberg,. 1906 

1906. Cervus elaphus scoticus Lonnberg, Arkiv. Zool. J, 9: 11. Glenquoich Forest, 
Inverness, Scotland. Range: England, Scotland, Ireland. 

Cervus elaphus macneilli Lydekker, 1909 

1909. Cervus cashmirianus macneilli Lydekker, P.Z.S. 588, pi. 69. Szechuan border of 


1910. Cervus canadensis wardi Lydekker, Abstr. P.Z.S. 38; 191 1, P.Z.S. iQio: 987. 

Szechuan border of Tibet. 

Cervus elaphus hispanicus Hilzheimer, 1909 

1909. Cervus elaphus hispanicus Hilzheimer, Archiv. fiir Rassen-und-Gesellschafts- 
Biol. 6: 313. Province of Huelva, between the Rio Odiel and the Guadal- 
quivir, Spain (Cabrera, 191 1). 

(?) 191 1. Cervus elaphus bolivari Cabrera, Bol. Soc. Esp. H.N. 11: 558. El Pardo, 
Madrid, Spain. 

Range: Spain and Portugal. 

Cervus elaphus kansuensis Pocock, 19 12 

1912. Cervus kansuensis Pocock, P.Z.S. 573. Thirty miles south-east of Taochow, 
Kansu, China. 

Cervus elaphus alashanicus Bobrinskii & Flerov, 1935 

1935. Cervus canadensis alashanicus Bobrinskii & Flerov, Arch. Mus. Zool. Moscou, /.■ 
29. Alashansk Range, South-Eastern Mongolia. 

Incertae sedis 

Elaphus minoratus Heude, 1892, Mem. H.N. Emp. Chin. 2: 113. No locality. 

Genus ELAPHURUS Milne-Edwards, 1866 

[ilne-Edwards, Ann. Sci. Nai 

I species : Elaphurus davidianus, page 37 1 

1866. Elaphurus Milne-Edwards, Ann. Sci. Nat. Zool. 5.- 382. Elaphurus davidianus 



Elaphurus davidianus Milne-Edwards, 1866 Mi-Iu, or Pere David's Deer 

Approximate distribution of species : the original range appears to have been the 
great alluvial plain of North-Eastern China, as far south as the Yangtze and Ch'ien- 
t'ang estuaries, but it became extinct in the wild state after the Shang Dynasty 
(Sowerby, 1949), and by the time Pere David sent his specimen home to Milne- 
Edwards the only survivors of this deer were those in the walled Imperial Hunting 
Park, south of Pekin. During the Boxer Rising in 1900 these deer escaped, and some 
were sent to Europe. The only deer to survive the rising in China were a few which 
were taken to Pekin itself where, by 191 1, only two specimens remained alive. By 
192 1 these had died. The only survivors today are the Duke of Bedford's herd at 
Woburn, founded by specimens sent to Europe in 1900, and a small herd at Whipsnade 
Zoological Park and a few specimens in the Bronx Zoo, New York, all derived from 
the Woburn herd. 

Elaphurus davidianus Milne-Edwards, 1866 

1866. Elaphurus davidianus Milne-Edwards, Ann. Sci. Nat. Zool. j.- 382. Imperial 

Hunting Park, Pekin, Chihli, China (captivity). 

1867. Cervus tarandoides David, Nouv. Arch. Mus. H.N. Paris, j, Bull.: 28. 

1933. Cervus [Rucervus) menziesianus Sowerby, China J. ig: 141. Near Anyang, 
Honan. (Based on fragments of antlers discovered in the course of archaeo- 
logical excavations.) 

Subfamily Odocoileinae 

As understood by Simpson (1945). Simpson makes each of the living Palaearctic 
genera the type of a Tribe. We exclude Hydropoles, as noted in the introduction to 
the Cervidae above. 

For characters of genera referred here, see Lydekker (1915). 

Genus CAPREOLUS Gray, 1821 

1775. Capreolus Frisch, Natur-System der vierfuss. Thiere, 3 (see page 2). 

182 1. Capreolus Gray, London Med. Repos. 15: 307. Cervus capreolus Linnaeus. 

1837. Caprea Ogilby, P.Z.S. 1836: 135. Cervus capreolus Linnaeus. 

I species: Capreolus capreolus, page 371 

Capreolus capreolus Linnaeus, 1758 Roe Deer 

Approximate distribution of species: widely distributed in the Palaearctic region 
(except in the extreme north and not occurring in North- Western India). 

(In detail: Britain, France, Spain and Portugal, Netherlands, Switzerland, Italy, 
Norway and Sweden, Germany, Poland, thence south-eastwards to Greece. Western 


l>.\L.\EARt:TIC: AND INDIAN MAMMALS 1 758-1946 

Russia (north approximately to Leningrad, with isolated populations in forests on 
Upper Don and in Crimea); Caucasus; Hissar and Alai Mountains, Tian-Shan 
Mountains, parts of Western Siberia (Lower Urals, basins of Middle Tobol and 
Ishim, whence it is spreading as far as Central Kazakstan and to the Irtish) ; Altai 
and Eastern Siberia, as far as Tatarsk Strait and Sea of Japan, north approximately 
to a line through Tomsk, Nishne-Ilimsk, northern tip of Lake Baikal, Southern 
Yakutia, etc. (Russian details condensed from Bobrinskii.) NLanchuria, Mongolia, 
\Vestern Chinese Turkestan, states of Chihli, Shansi, Shensi, Kansu, Szechuan in 
China. Persia, and Asia Minor, to Northern Iraq.) 

Capreolus capreolus capreolus Linnaeus, 1758 

1758. Cenus capreolus Linnaeus, Syst. Nat. loth ed. /.• 68. Sweden. 

1792. Cervus capreolus alhus Kerr, Anim. Kingd. 302. Tranche Comte, France. 

1830. Capreolus dorcas Burnett, Quart. J. Sci. Lit. Art. iSsg, 2: 353, nom. nud. 

1832. Capreolus vulgaris Fitzinger, Beitr. Landesk. Osterreichs, /.• 317. 

1843. Capreolus capraea Gray, List. Spec. Mamm. B.M. 176. Renaming of capreolus. 

1845. Cervus capreolus plumbeus Reichenbach, Naturg. Saugeth. 3, pi. 3 bis, fig. 53. 


1846. Capreolus europaeus Sundevall, K. Svenska Vetensk. Akad. Handl. 1844: 184. 
1874. Capreolus vulgaris niger Fitzinger, S.B. Akad. Wiss. \Vien, yo, i : 247. Germany. 
1874. Capreolus vulgaris varius Fitzinger, loc. cit. Germany. 

iqo7. Capreolus transsylvanicus Matschie, Weidwerk in Wort u. Bild, 16: 224. Bana, 

iQio. Capreolus eapreolui balticus Matschie, Weidwerk in Wort u. Bild, ig: 263. 

Wichcrtshof, East Prussia, i April, 1910.) 
igio. Capreolus capreolus albicus Matschie, loc. r//. Jesziorki, near Lissa, Poland. 
iQio. Capreolus capreolus rhenanus Matschie, loc. cit. Rouflach, Haute-Rhin, France. 
iqio. Capreolus capreolus Ihotti Lonnberg, Ann. Mag. N.H. 6: 297. (September, 1910.) 

Arndilly, Craig EUachie, Morayshire, Scotland. (Not Aberfeldy, as stated 

by Lonnberg.) 
191 o. Capreolus capreolus canus Miller, Ann. Mag. N.H. 6: 460. f November, 1910.) 

Quintanar de la Sierra, Burgos, Spain. 
iqi2. Capreolus capreolus ivarthae Matschie, Dtsch. Jagerztg. 5-9.- 801. Dombrowa, east 

of Bcuthcn, Poland. 
1013. Cervus i Capreolus) capreolus eistaumcus Matschie, VerofT. Inst. Jagdk. Neudamm, 

2: 139. Dunnwald, north of Cologne, Germany. 
iqi3. Cervus [Capreolus) capreolus transvosagicus Matschie, loc. cit. Staufcn, in the 

Vosgcs, Eastern France. 
i()i6. Capreolus capreolus decorus Cabrera, Bol. Soc. Esp. H.N. 16: 175. El Vierzo, 

Province of Leon, Spain. (March, 1916.) 
n)iG. Capreolus capreolus armenius Blackler, Ann. Mag. N.H. 18: 78. Sumela, near 

Trebizond, Asia Minor. (July, 1916.) 
K)i6. Capreolui capreolus joffrei Blackler, Ann. .Mag. N.H. 18: 79. Ferrieres, Paris, 

i()i6. Capreolus zedlitzi Matschie, S.B. Gcs. Naturf. Fr. Berlin, 272. Slonim, Poland. 

'December, 1916.) 
iq23. Capreolus co.xi Chceam^in & Hinton, Ann. Mag. .N.H. 12: 608. Zakho, Northern 




1925. Capreolus capreolus italicus Festa, Boll. Mus. Zool. Anat. Comp. Torino, 40, 37: 

I. Castelporziano, Central Italy. 
1925. Capreolus capreolus grandisBolkay , Novit. Mus. Sarajevo, /.• 14. Neighbourhood 

of Sarajevo, Yugoslavia. 
■933- Capreolus capreolus grandis morpha baleni Martino, O sar-planinskom srndacu, 2 

(of reprint). Shar-Planina, borders of Albania and Yugoslavia. 
1936. Capreolus capreolus whittalli Barclay, Ann. Mag. N.H. ly: 405. Near Alemdagh, 

15 miles from Moda, Istanbul, Turkey. 

Range: Europe, including England, Scotland, \Vales, Russia, Asia Minor, Persia. 

Capreolus capreolus pygargus Pallas, 1771 

1 77 1. Cervus pygargus Pallas, Reise Russl. /.■ 453. River Sok, Samara district, Volga, 

1906. Capreolus tianschanicus Satunin, Zool. Anz. jo." 527. Kuldja, Chinese Tian Shan 

1909. Capreolus pygargus firghanicus Rasewig, Semja ochoton. Moskva, igog: 160. 

Fergana district, Russian Turkestan. [N.V.) 

Range: Central Asia (Altai and Tian Shan Mountains), westwards to the Urals and 
the Volga; northerly and easterly distribution uncertain. 

Capreolus capreolus bedfordi Thomas, 1908 

1908. Capreolus bedfordi Thomas, Abstr. P.Z.S. 32; 1908, P.Z.S. 645. Mt. Chao- 
Cheng-Shan, 100 miles west-north-west of Taiyuenfu, Shansi, China. 

1889. Cervus pygargus mantschuricus Noack, Humboldt, 8: 9. Eastern Manchuria. Not 
mantchuricus Swinhoe, 1864. 

1911. Capreolus melanotis Miller, Proc. Biol. See. Washington, 24: 231. Thirty miles 
east of Ching-yang-fu, Kansu, China. 

1935. Capreolus capreolus ochracea Barclay, Ann. Mag. N.H. /j.- 627. Korea. 

Range: Szechuan, Kansu, Shansi, Chihli, Manchuria, Korea, Northern Mongolia, 
South-Eastern Siberia (Amur-Ussuri region). Quelpart I. 

Genus ALCES Gray, 1821 

1775. Alee Frisch, Natur-System der Vierf. Thiere, 3. 

1 82 1. Alces Gray, London Med. Repos. 75 .• 307. Cervus alces Linnaeus. (Opinion 91 
of the International Commission on Zoological Nomenclature says that the 
European Elk should be called Alces Gray, 1821, with Cervus alces Linnaeus 
as type species.) 

1841. Alcelaphus Gloger, Handb. Naturges. 143. (Substitute for Alces.) 

1902. Paralces ]. Allen, Bull. Amer. Mus. N.H. 16: 160. (Substitute for Alces.) 

I species: Alces alces, page 373 

Alces alces Linnaeus, 1 758 Moose, or Elk (in European sense) 

Approximate distribution of species: Norway, Sweden, East Prussia, thence east- 
wards across Russia and Siberia (not including Kamtchatka) ; Manchuria, Mongolia. 
Northern North America. 


palaearc:tic and Indian mammals 1758-1946 

For European details, see under A. a. alces. In U.S.S.R., "in East Europe and 
Trans-Lena Siberia the northern hmit in general coincides with that of full-grown 
forest, but in West and to some extent in Central Siberia it comes a long way south 
in a wide, shallow arc roughly to the Goth parallel. . . . The southern limit in the 
U.S.S.R. runs roughly along the southern border of \Vhite Russia, proceeds to 
Ryazan, dips far south in a long tongue, returns north almost to Gorki, turns south 
again but in a wide tongue along the right side of the Volga, approximately 
to 52^ 50', continues along the Kama and then the Lower Belaya, dips down 
along the Southern Urals roughly to IvLagnitogorsk and returns north almost to 

South of this line, particularly in area between the Don and Volga, elks make 
fairlv long incursions. They also occur isolated in the Buzuluk pine forest (between 
Kuibuishev and Chkalov). In Siberia the limit runs roughly from Sverdlovsk to 
Tomsk, roughly coinciding with the southern limit of the continuous taiga, reaches 
the Yenesei a little below Krasnoiarsk, skirts round the east of the Minussinsk 
steppes, takes in the Eastern Altai, and passes out into Northern Mongolia. Beyond 
the Yenesei the elk fails to occur in the steppe parts of Transbaikalia and in the 
extreme south of Ussuri region, and is not found in Sakhalin" i Bobrinskii).) 

Alces alces alces Linnaeus, 1758 

1758. Cervus alces Linnaeus, Syst. Nat. loth ed. /.■ 66. Sweden. 

1827. Cervus coronatus Lesson, Man. Manim. 356. 

1830. Alces europaeus Burnett, Quart. J. Sci. Lit. Art. i82g, 2: 353, twm. nud. 

1837. Alces machlis Ogilby, P.Z.S. 1836: 135. Renaming oi alces. 

1841. Alcelaphus alee Gloger, Handb. Naturgesch. /.• 143. 

1842. Alces antiqiwrum Ruppell, Mus. Senckenb. jj.- 183. Renaming oi alces. 

1843. Alces palmalus Gray, List Mamm. Coll. B.M. 182. 

i860. Alces jubata Fitzinger, Nat. Saugeth. .}: 86 [N.V.) Renaming oi alces. 

icjio. Alces machlis typicus Ward, Rec. Big Game, cd. 6, 99. 

191 3. Alces machlis uralensis Matschie, Veroff. Inst. Jagdk. Neudamm, :?.- 155. District 

of Samara, Russia. 
191 5. Alces machlis tymensis Zukowsky, Arch. Naturgesch. Berlin, ScjA, 9; 42. River 

Tym, Siberia. 
1915. Alces machlis angusticephalus Zukowsky, loc. cit. 44. Yeneseisk District, 

Range: Scandinavia, Lapland, Finland, Baltic States, East Prussia, Poland, Russia 
and Siberia as far as Yenesei River and Lake Baikal. 

.■\lces alces cameloides Milne-Edwards, 1867 

i8r)7. Cervus cameloides .Milne-Edwards, Ann. Sci. Nat. Zool. y: 377. Probably from 

1902. Alces hedfordiae Lydekker, P.Z.S. icjo2, i: 109. Siberia, 
iqio. Alces pfi.zenmayeri Zukowsky, Wild und Hund, 16: 807. River .A.ldan, Xorth- 

Eastern Siberia, 
iqii. Alces machlis yakutskensis Millais, The Field, London, 118: 113. River Aldan, 

North-Eastern Siberia (based on same material as pfizenmayeri) . 
Range: Siberia, east of the Yenesei, Mongolia, Manchuria. 



Genus RANGIFER H. Smith, 1827 

1775. Rangifer Frisch, Natur-System der vierfuss. Thiere, 3. 

1827. Rangifer H. Smith, Griffith's Cuvier Anim. Kingd., Mamm. Syn. 304. Cenms 

tarandus Linnaeus. 
(Opinion 91 of the International Commission on Zoological Nomenclature 

states that Rangifer should date from H. Smith, 1827, with Cervus tarandus 

Linnaeus as type species.) 
1827. Tarandus Billberg, Syn. Faun. Scand. /.■ 22. Tarandus lapponum Billberg = 

Cervus tarandus Linnaeus. 
1845. Achlis Reichenbach, Naturges. Saugeth. 2- 12. Alternative for Tarandus. 

See Jacobi, A., 1931, Das Rentier, ^ool. Anzeiger, g6 (Erganzungsbd) : 1-264. 
Flerov, C. C., 1933, Review of the Palaearctic Reindeer or Caribou, J. Mammal, 
14: 328. 

I species: Rangifer tarandus, page 375 

Rangifer tarandus Linnaeus, 1758 Reindeer, Caribou 

Approximate distribution of species : Arctic regions of Old and New Worlds, local 
distribution modified by human agency. Norway, Spitzbergen, Finland, Arctic 
regions of Russia, Arctic Siberia, east to Kamtchatka and Sakhalin, south to Mon- 
golia and nearly to Chinese border in the east (Bobrinskii). Arctic regions of North 
America, Greenland included. 

Rangifer tarandus tarandus Linnaeus, 1758 

1758. Cervus tarandus Linnaeus, Syst. Nat. loth ed. /.• 67. Alpine region of Swedish 

1788. Cervus tarandus rangifer Gmelin, Syst. Nat. 13th ed. /.• 177. 
1827. Tarandus lapponum Billberg, Synops. Faun. Scand. /.■ 22. Renaming of 

1842. Tarandus borealis Ruppell, Mus. Senckenb. ^.- 183. Renaming oi tarandus. 
1852. Tarandus furcifer Baird, Rep. Comm. Patents, 1851, 2, Agric. 109. [M.V.) 

Renaming of tarandus. 
1898. Rangifer tarandus typicus Lydekker, Deer of all Lands, 38. 
1902. Rangifer tarandus var. cilindricornis Camerano, Mem. R. Accad. Torino, j/.- 

167. Renaming of teranrfaj. 
1909. Rangifer tarandus fennicus Lonnberg, Arkiv. Zool. 6, 4: 10. Torne, Lappmark, 

1936. Rangifer tarandus silvicola Hilzheimer, Z. Sauget. //.■ 155. Olenez district, 


Range: Scandinavia, to Russia. 

Rangifer tarandus platyrhynchus Vrolik, 1829 

1829. Cervus {Tarandus) platyrhynchus Vrolik, Nieuwe Verh. Konink. ^ederl. Inst. 

Eerste Klasse, 2: 160. Spitzbergen. 
1862. Cervus tarandus forma spetsbergensis Andersen, Ofvers. Vek. Akad. Forhandl. ig: 

457. Spitzbergen. 


P.\LAE.\RC:TIC and INDIAN MAMMALS 1 758-1946 

Rangifer tarandus platvrhynchus [contd.] 

1866. Rangifer arcticiis var. spilzbergensis Murray, Gcogr. Distrib. Mamm. 334 (des- 
cribed on p. 155). Spitzbergen. 

Rangifer tarandus sibiricus Murray, 1866 

i86(3. Rarigiftr tarandus sihiricus Murray, Geogr. Distrib. Mamm. 334 (described on 

p. i'it). Siberia, eastward of the River Lena, [sibiricus Schreber, 1784, is not 

a valid name. The word is used to indicate the provenance of the particular 

reindeer illustrated in pi. 284C of Theil 5.) 
191 r,. Tarandus rangifer lenensis Millais, the Big Game of Asia and North America, 

2 If) (The Guri at Home and Abroad, 4). Delta of River Lena. 
1915. Tarandus rangifer clmkchensis Millais, loc. cit. 220. Delta of Ri\er Lena (a 

domesticated form). 
1915. Tarandus rangifer yakutskensis Millais, loc. cit. 222. Yakutsk (a domesticated 


1 93 1. Rangifer arcticus asiaticus ]i^cob\, Zool. .\nz. ry6" (Erganzungsbdi : 85. Kolyma 

River, North-Eastern Siberia. (Renaming o( sibiricus Murray.) (The type 

was a tame animal.) 
1936. Rangifer tarandus transuralensis Hilzheimcr, Z. Sauget. //; 155. Konda River, 

Western Siberia. 
Range: Siberian and Eastern European tundra zone; Novosibirskie Islands, Arctic 

Rangifer tarandus pearsoni Lydekkcr, 1903 

1903. Rangifer tarandus pearsoni Lydekker, P.Z.S. icjo2, 2: 361. Lsland of Novaya 

Rangifer tarandus phylarchus Hollister, 191 2 

1912. Rangifer phylarchus Holhster, Smiths. Misc. Coll. j6', 35: 6. South-Eastern 
Kamtchatka. Range includes coast of Okhotsk Sea and Amurland. 

Ran-qifer tarandus angustirostris Flerov, 1932 

1932. Rangifer angustirostris Flerov, Trav. Comin. Rep. S.S. Lakoute, 4: 8. Bargusin 

Mountains, north-eastern coast of Lake Baikal. 

Rangifer tarandus valentinae Flerov, 1933 

1933. Rangifer tarandus valentinae Flerov, J. NLamm. 14: 336. Head of Chulyshman 

River, North-Eastern Altai, Siberia. 
(?) 191 5. Tarandus rangifer buskensis Millais, The Big Game of Asia- and North 

America, 222"("The Gun at Home and Abroad, 4). Busk Mountains, near 

Semipalatinsk, Siberia. 
Range: forest zone of Siberia, south to Northern Mongolia and Altai Mountains. 

Rangifer tarandus setoni Flerov, 1933 

1933. Rangifer tarandus setoni Flerov, J. .NLamm. 14: 337. Sakhalin Island, Eastern 




Genera: Addax, page 384 Hemitragus, page 403 

Ammotragus, page 409 Naemorhedus, page 401 

Antilope, page 386 Oryx, page 385 

Bison, page 382 Ovis, page 411 

Bos, page 379 Pantholops, page 395 

Boselaphus, page 379 Procapra, page 387 

Bubalus, page 383 Pseudois, page 410 

Budorcas, page 396 Rupicapra, page 397 

Capra, page 404 Saiga, page 395 

Capricornis, page 399 Tetracerus, page 378 
Gazelle, page 388 

Alcelaphus Blainville, 1816, £«//. 6'oc. Philom. Paris, 75 [^ Bubalis Frisch, 1775, 
J^atur-Sjslem der vierfuss. Thieve, 2 — unavailable, see p. 2) ; Alcelaphus buselaphus 
buselaphus (Antilope buselaphus Pallas, 1 766, jA/wf . ,^00/. 7 — type locality probably 
Morocco) formerly ranged across North Africa from Morocco to Egypt, but became 
extinct some time in the 1920's. Other races of the species buselaphus occur in Africa, 
from Senegal to Somaliland, and Tanganyika. For details and for a full synonymy 
of the typical race, see G. Allen, 1939, Checklist of African Mammals, 470. 

This family is very difficult to classify and no two authors agree on the various 
subfamilies or minor divisions, some of which seem to be indefinable and un- 
convincing. The characters of most of the species and genera are to be found in 
Lydekker, but his key to subfamilies, spread as it is over three different volumes, is far 
from clear. 

Blanford (1891, 482) gives a key to the genera inhabiting India. But, in part, this 
may not hold good for species or genera extralimital to India. 

Of the genera here listed, Addax and Ammotragus are solely African. 

An extremely interesting and instructive paper on this family is Pocock, 191 1, 
On the specialized cutaneous glands of ruminants, P.^.S. igio: 840. 

Simpson (1945, 270-272) discusses the grouping of the family in some detail, and 
his classification is followed here. According to his list, with some slight generic 
modification, the living genera now under discussion are classified as follows : 

Subfamily: Bovinae 

Tribe: Boselaphini 

Boselaphus, Tetracerus 
Tribe : Bovini 

Bos, Bison, Bubalus 

Subfamily: Hippotraginae 

Tribe: Hippotragini 

Addax, Oryx 

(Tribe: Alcelaphini 


* Now extinct in the Palaearctic region. 



Subfamily: Antilopinae 

Tribe: Antilopini 

Ant Hope, Gazdla, Procapra 

Subfamily: Caprinae 

Tribe : Saigini 

Saiga, Pantholnps 

Tribe: Rupicaprini 

Capricornis, Naemorltediis, Riipicapra 

Tribe : Ovibovini 

Tribe: C'aprini 

Hemitragus, Capra, Pseudois, Ammotragus, Ovis 

Subfamily B o v i n a e 

(As understood by Simpson, 1945) 

Many authors prefer to refer Boselaphus and Tiiracerus to a distinct subfamily 
Bosclaphinae, and restrict the Bovinae to Bos, as here understood. Bison and Biibahs. 

Genus TETRACERUS Leach, 1825 

1825. Tctracerus Leach, Trans. Linn. Soc. London, 14: 524. Antilope chickara Hard- 
wicke = Cerophorus quadricornis Blainvillc. 

1827. Tctraceros "Qrodkes, Brookesean Museum (2nd ed.), 8. For Tetracerus. 

I species: Tetracerus quadricornis, page 378 

Tetracerus quadricornis Blainville, 1816 Four-horned Antelope; Ghousingha 

Approximate distribution of species: Peninsula of India; from Madras and Eastern 
Ghats north at least to Kathiawar and Central Pnix'inces. 

Tetracerus quADRicoRNis Blainville, 1816 

1816. Cerophorus (Cervicapra) quadricornis Blain\illc, Bull. Soc. Philom. Paris, 75 and 

78. Plains of Peninsular India. 
1825. Antilope chickara Hardwicke, Trans. Linn. Soc. London, i^: 520, pis. 15, 16. 

Western Provinces of Bengal, Bihar and Orissa, India. 

1828. Tetraceros striatocornis Brookes, Cat. Mus. 64. No locality. 

1836. Antilope tetracornis Hodgson, J. Asiat. Soc. Bengal, 4: 525. No locality. 

183C). Antilope sub-^-cornutus Elliot, Madras J. Lit. 10: 225. Southern Mahratta 

country, India. 
1843. Tetracerus subquadricornis Gray, List Mamm. B.M. 159. 
1847. Tetracerus subquadncornutus Hodgson, Calcutta J.N.H. 8: 89. Emendation of 



1847. Tetracerus iodes Hodgson, Calcutta J.N.H. 8: 90. "Saul forests beneath the 

1847. Tetracerus paccerois Hodgson, loc. cit., same locality. 
1895. Tetraceros quadricornis typicus Sclater & Thomas, Book of Antelopes, /.■ 215. 

Cerophorus Blainville, 1816, Bull. Soc. Philom. Paris, 74, is really equivalent to 
"Bovidae" since it includes all ruminants "qui sont toujours la tete armee", not 
counting the giraffe, as opposed to camels, and deer (where the head armature is 

Genus BOSELAPHUS Blainville, 1816 

1816. Boselaphus Blainville, Bull. Soc. Philom. Paris, 75. Antilope picta Pallas = 

Antilope tragocamelus Pallas. . 
1827. Portax H. Smith, Griffith's Cuvier Anim. Kingd. j.- 366. Damalis risia Smith = 

Antilope tragocamelus Pallas. 
1851. Bosephalus Horsfield, Cat. Mamm. Mus. E. Ind. Co. i6g. Error for Boselaphus. 

I species: Boselaphus tragocamelus, page 379 

Boselaphus tragocamelus Pallas, 1766 Nilgai; Blue Bull 

Approximate distribution of species: Peninsular India, from the base of the 
Himalayas to Mysore, in Eastern Punjab, Gujerat, North-^Vest Provinces and parts 
of Bombay. (Not in Eastern Bengal, or east of that, and not on the Malabar coast.) 
(Dunbar Brander, 1927.) 

Boselaphus tragocamelus Pallas, 1 766 

1766. Antilope tragocamelus Pallas, Misc. Zool. 5. Plains of Peninsular India. 

1777. Antilope albipes Erxleben, Syst. Regn. Anim. 280. India. 

1777. Antilope picta Pallas, Spicil. Zool. 12: 14. India. 

1827. Damalis risia H. Smith, Griffith's Cuvier Anim. Kingd. 4: 363. Substitute for 

1837. Tragelaphus hippelaphus Ogilby, P.Z.S. iS^G: 138. Substitute {or picta. 

1846. Portax tragelaphus Sundevall, K. Svenska Vetensk. Akad. Handl. 1844: 198. 

Corrected to tragocamelus, 184^: 323. 

Genus BOS Linnaeus, 1758 

1758. Bos I.innaeUs, Syst. Nat. loth ed. /.• 71. Bos taurus Linnaeus (Domestic Cattle). 

1814. Taurus Rafinesque, Princip. Somiol., 30. Renaming of Bos Linnaeus. 

1827. Urus H. Smith, Griffith's Cuvier Anim. Kingd. 4: 417. Urus scoticus Smith 

(white Park Cattle). 
1837. Bibos Hodgson, J. Asiat. Soc. Bengal, 6: 499. Bibos subhemachalus Hodgson = 

Bos gaurus H. Smith. Valid as a subgenus. 
1843. Poephagus Gray, List Mamm. B.M. 153. Bos grunniens Linnaeus. Valid as a 


1847. Gaveus Hodgson, J. Asiat. Soc. Bengal, 16: 705. Bos frontalis Lambert. 
1901. Gauribos Heude, Mem. H.N. Emp. Chin. 5, 1:3. Gauribos laosiensis Heude. 



Bos [contd.] 

1 90 1. Urihos Hcudc, Mem. H.N. Emp. Chin. 5, i : f,. Unhos platyceros Hcude. 
1901. Biihtilihoi Hcudc, Mem. H.N. Emp. Chin. 5, i : (>. Buhalihos annamilicus Hcude. 
1940. Novibos Coolidge, Mem. Mus. Comp. Zool. Harvard, 5^.- 425. Bos {Bibos) 
sauveli Urbain. 

4 species: Bos hanteno, page 381 
Bos gaunis, page 38 1 
Bos grunniens, page 382 
Bos samvli, page 382 

The type species is domestic. The generic divisions for the various kinds of wild 
o.xen Hving today are not very clear. As many as eight genera have been recognized : 
Bos. Btbos, Novibos, Po'cphagus, Bison, Anoa, Bubalus and Syncerus (the last three com- 
prise the buffaloes). Those who regard all eight as valid genera restrict Bos to 
domestic oxen and their extinct allies. Lydekker (1913) referred all oxen to one 
genus Bos, and recognized four extra subgenera, Bibos, Po'cphagus, Bison and Bubalus 
(the last containing all buflaloes). 

Other authors compromise between these two extremes, and Simpson (1945) 
recognizes all .save Novibos and Po'cphagus. The latter he refers to Bos, though it would 
seem that it is more worthy of generic distinction than Bibos. Novibos, which Coolidge 
erected (on the basis of one specimen) in 1940 for the Kouprey of Cambodia seems 
doubtfully valid. 

If there is to be generic distinction between the living ox-like Bovinae, then it 
should be between the buffaloes, for which the prior name is Anoa, and the remainder. 
This view appears to be supported, too, by Pilgrim's work on living and fossil 
Bovidae. It is adopted in this list, but although Pocock (1918) regarded Anoa and 
Bubalus as congeneric, we do not feel it is advisable to use Anoa as the generic name 
for the Indian buffalo, so we follow those authors who restrict that name to the 
dwarf species from Celebes and Philippines. Also Bison, universally admitted by 
American authors and much the most distinct of the groups which perhaps should 
be referred to the genus Bos, is tentatively retained. 

Subgenus BIBOS Hodgson, 1837 

The first name in the subgenus is Bos frontalis Lambert, 1804, Trans. Linn. Soc. 
London, j: 57, which was based on a specimen of the domestic Gayal, probably from 
North-Eastern Chittagong. Lydekker (191 3) regarded the next name. Bos sylhctanus 
F. Cuvier, in GcofTroy & Cuvier, 1824, H..'N'. Mamm. j, 42: Jungly-Gau, 2, and 
pi. 418, from Sylhet, Assam, as referring also to the Gayal. But Cuvier's plate and 
description seem to be more applicable to the Gaur than the Gayal. However, since 
there is room for doubt, and since Bos gaurus Smith, 1827, is the name by which 
the Gaur is now generally known, it is proposed as a matter of convenience to 
accept Lydckkcr's interpretation rather than revive the name sylhctanus for the 



Bos gaurus H. Smith, 1827 Gaur (or Indian "Bison") 

Approximate distribution of species: Malay States, Indo-China, Burma, Assam, 
Nepal, Peninsular India in forest areas, south to Travancore. 

Bos GAURUS GAURUS H. Smith, 1827 

1827. Bos gaurus H. Smith, Griffith's Cu\'ier Anim. Kingd. 4: 399. Mainpat, in the 

Sarguja Tributary States, India, in approximately 23° N., 83' E. (Harper, 

1827. Bos gour Hardwicke, Zool. J. 5; 231. "District of Ramgurh and tableland of 

Sirgoojas," India. 
1837. Bibos subhemachalus Hodgson, J. Asiat. Soc. Bengal, 6: 499. Saul Forest, Nepal. 
1837. Bos cavifrons Hodgson, J. Asiat. Soc. Bengal, 6: 747. Substitute for subhemachalus. 
184(5. Bos gaur Sundevall, K. Svenska Vet. Akad. Handl. 1844: 201. Substitute for 

1851. Bos asseel Horsfield, Cat. Mamm. Mus. E. Ind. Co. 181. 
Range: Peninsular India to Rajputana, Nepal, Bhutan Duars, Assam. 

Bos GAURUS READEi Lydckkcr, 1903 

1903. Bos gaurus readei Lydekker, Zoologist, y: 266. Myitkyina, Upper Burma. 
Range: Burma, Tenasserim, Cochin-China. 

Bos banteng Wagner, 1844 Banteng; Tsaine 

Approximate distribution of species: Burma, Siam, Indo-China, Malay States, 
Java and Borneo. 

(Bos BANTENG BANTENG Wagner, 1844. Extralimital) 

1844. Bos banteng W'agner, in Schreb. Saugeth. Suppl. 4: 517. Java. 

1845. Bos sondaicus Schlegel & Miiller, in Temminck, Verh. Nat. Gesch. Ned. 

Overz. Bezitt. Zool. Mamm. 197. Java. 

1846. Bos banting Sundevall, K. Svenska Vetensk. Akad. Handl. 1844: 152. Java. 

Bos BANTENG BiRMANicus Lydckker, 1898 
1898. Bos sondaicus hirmanicus Lydekker, P.Z.S. 277. Burma. 
(?) 1909. Bos sondaicus porteri Lydekker, P.Z.S. 66g. Siam. 
Range: Burma, Siam, Indo-China. 

The following names were given by Heude to Indo-Chinese specimens of the s,\ih- 
genus Bibos. One of them may be valid if an Indo-Chinese race proves separable, and 
there is always the chance that one of them may prove to antedate B. sauveli, below. 

1901. Gauribos laosiensis Heude, Mem. H.N. Emp. Chin. 5, 1:3. Laos, Tonkin. 

1 90 1. Gauribos brachyrhinus Heude, loc. cit. 4. Pursat, Cambodia. 

1 90 1. Gauribos sylvanus Heude, loc. cit. 4. Mois, Tonkin. 

1 90 1. Gauribos rnekongensis Heude, loc. cit. 5. Kratie, Mekon Valley. 

1901. Uribos platyceros Heude, loc. cit. Range dividing Tourane Bay from the rivers 

of Hue. 
1901. Bubalibos annamiticus Heude, loc. cit. 6. Annam. 
1901. Bos (?) leptoceros Heude, loc. cit. 7. Kampot, on coast of Gulf of Siam. 



1901. Bibos discolor Heude, loc. cit. 8. No locality. 
'901. Bihos longicornis Hcude, loc. cit. 9. No locality. 
190'. Bibos {?) fusicornis Heude, loc. cit. g. Tonkin. 

Bos sauveli Urbain, 1937 Knuprey, or Cambodian Forest Ox 

.Approximate distribution of species: Cambodia (Indo-China). 

Bos s.^uvELi Urbain, 1937 

1937. Bos [Bibos) sauveli Urbain, Bull. Soc. Zool. France, 62: 307. Near the village 
of Tchep, North Cambodia (Urbain, 1939). 

On this species, see the monograph by Coolidge, 1941, Mem. Mtis. Comp. ^ool. 
Harvard. ^4: 421-531, where the animal's affinities are fiilly discussed and illustrated. 

F. Edmond Blanc, 1947, .K contribution to the knowledge of the Cambodian Wild 
Ox or Kouprey, J. Mamm. 28: 245-248, suggests this species is a hybrid between the 
Banteng and either the Gaur, Water Buffalo or domestic cattle. 

Subgenus POEPHAGUS Gray, 1843 

Bos grunniens Linnaeus, 1766 Vak 

Approximate distribution of species: Tibet, Kansu, Ladak. The domesticated 
form, variously in the high plateaux and mountains of Central Asia. (Prater states 
that within Indian limits proper, yak only occur in the Chang Chen Mo Valley, in 
Ladak. They sometimes stray into the Sutlej Valley and into some of the passes in 
Eastern Kumaon.) 


1 766. Bos grunniens Linnaeus, Syst. Nat. r2th ed. /.■ 99. 'Tn Asia boreali." The species 

was based on the domesticated breed. 
181 1. Bos poephagus Pallas, Zoogr. Ross. Asiat. /.• 248, pi. 22. Renaming oi grunniens. 
1833. Poephagus gruniens Gray, List Mamm. B.^L 153. 

Bos GRUNNIENS MUTUs Przcwalski, 1883 (Wild Yak) 

1883. Poephagus mutus Przewalski, Third Journey in C. .Asia, igi. Alpine region of the 
western part of the Nan Shan (appro.ximately 39^20' N., 95° E.), between 
the Anembar-Ula on the west and the Humboldt Range on the east. 
Northern Kansu, China. (Harper, 1940.) 

Genus BISON H. Smith, 1827 

1827. Bi<<on H. Smith, Griffith's Cuvier Anim. Kingd. j.- 373. Bos bison Linnaeus 
(Opinion 91 of the International Commission on Zoological Nomenclature), 
the North American Bison. 

1844. Bonasus Wagner, Schreb. Saugcth. Suppl. 4: 515. Bos (Bison) bison Smith = 
Bos bonasus Linnaeus. 

I species in the area covered by this list : Bison bonasus, page 383 

There are two existing species, one of which is American. 



Bison bonasus Linnaeus, 1758 European Bison (Wisent) 

Approximate distribution of species: Lithuania, and till recently in the Caucasus. 
For details of the history in both places see J. Soc. Pres. Fauna Emp. 1949, pt. 59. 

Bison bonasus bonasus Linnaeus, 1 758 

1758. Bos bonasus hirxaaLtvii,, Syst. Nat. lothed. /.• 71. Probably Bialowieza, Lithuania 

(Lydekker, 1913). 
1785. Bos urus Boddaert, Elench. Anim. /.• 151. 
1827. Bos bison H. Smith, Griffith's Cuvier Anim. Kingd. 4: 398. Poland. Not of 

Linnaeus, 1758. 
1827. Urus nostras Bojanus, Nova Acta Leop. Carol, /j; 413. Bialowieza, Lithuania. 
1849. Bison europaeus Owen, P.Z.S. 1848: 126. Alternative for bonasus. 

The Lithuanian Bison became extinct at Bialowieza in 1 921, so far as the original 
free-living stock is concerned, but there were at that time some 45 specimens in 
zoological gardens and parks in Europe. By 1949 this number had been increased to 
119 pure-blooded specimens, including those which have been re-established in a 
reservation in the Bialowieza forest. 

Bison bonasus caucasius Greve, 1906 

igo6. Bison bonasus var. caucasia Greve, Zool. Beob. 4j: 270. Caucasus Mountains, 
South-Eastern Russia. Became extinct about 1925. 

Genus BUBALUS H. Smith, 1827 

1775. Bubalus Frisch, Natur-System der vierfiiss. Thiere, i (see page 2). 

1827. Bubalus H. Smith, Griffith's Cuvier Anim. Kingd. 5.' 2,T^-Bos bubalus Gmelin = 

Bos bubalis Linnaeus. 
1865. Bujfelus Rotimeyer, Verh. Naturf. Ges. Basel, 4: 334. Buffelus indicus Rutimeyer 

= Bos bubalis Linnaeus. 

I species : Bubalus bubalis, page 383 

Bubalus bubalis Linnaeus, 1758 Indian Buffalo; Water Buffalo; Arna 

Appro.ximate distribution of species: domesticated variously, including to Egypt. 
In India, Prater states "the grass jungles of the Nepal Terai and the plains of the 
Ganges and Brahmaputra in Assam; a few herds survive in parts of Orissa, adjoining 
the Raipur district of the Central Provinces and in the south-eastern districts of the 
Central Provinces". ? Ceylon (probably feral). Indo-China. A race is named from 

Bubalus bubalis bubalis Linnaeus, 1758 

1758. Bos bubalis Linnaeus, Syst. Nat. loth ed. /.• 72. Rome, Italy. (Thomas, 191 1, 
P.Z.S. 154. Linnaeus' description was based on a domesticated form.) 

1788. Bos bubalus Gmelin, Syst. Nat. /.■ 206. Asia. 

1792. Bos arnee Kerr, Anim. Kingd. 336. "India, north from Bengal," restricted to 
Kuch Bihar. (Harper, 1940.) 

1821. Bos bujfelus Blumenbach, Handb. Naturges. ed. 10, iig. Asia. 




1827. Bos ami H. Smith, Griffith's Cu\icr Anini. Kingd. 4: 388. Central Bengal. 

1841. Bubalus arna Hodgson, J. Asiat. Soc. Bengal, lo: 469. 

(?) 1842, Buhalus arna var. macrocerus Hodgson, J. Asiat. Soc. Bengal, 10: gi2, 7wm. 
nud. Assam (Lydekker, 1913, Cat. Ung. /.• 45). 

1842. Buhalus arna var. speirocerus Hodgson, loc. cit., nam. nud. 

1865. Bujfdus indkus Rutimeyer, Verh. Naturf. Ges. Basel, 4: 334. India. 
1865. Bujftlus indiais var. italica Rutimeyer. loc. cit. Italy (domestic). 
(?) 191 2. Buhalus huhalus septentrionalis Matschie, Deutsche Jager-Zeitung, 59.- 103. 
"Kuckri-Muckri," \orth-\\'estcrn India. 

Bubalus bub.-\lis fulvus Blanford, 1891 

1891. Bos huhalus \-a.r. fulvus Blanford, Fauna Brit. India, Mamm. 492. Mishmi Hills, 
Upper Assam. 

Subfamily H i p p o t r a g i n a e 

(As understood by Simpson, 1945) 

This subfamily, admitted by Simpson (p. 272) to be less surely natural than the 
other subfamilies he recognized, has something of the appearance of a wastepaper 
basket. We have our doubts whether it could be defined. The genus Alcelaphus has 
usually been regarded as belonging to a subfamily Alcclaphinae distinct from the 

Genus ADDAX Laurillard, 1841 

1 815. Addax Rafinescjue, Analyse de la Nature, 56, nom. nud. 

1841. Addax Laurillard, in d'Orbigny's Diet. Univ. Hist. Nat. /.■ 619. Antilope 
suturosa Otto = Cerophorus nasomaculata Blainville. 

(For date of publication, see Sherborn & Palmer, 1899, Ann. Mag. N.Il. 3: 350.) 

I species: Addax nasnmaculatus, page 384 

Addax nasomaculatus Blainville, 181 6 Addax 

Approximate distribution of species: Senegambia (doubtful) and Rio de Oro, 
Algerian Sahara and Southern Tunisia south to about 15° N., and east to the Anglo- 
Egyptian Sudan (Dongola, Darfur, Kordofan). Extinct in Egypt about the year 
1900 (Flower, 11)32). 

Adda.x nasomaculatus Blain\illc, 1816 

1816. Cernfiliorus (Gazilla) nasomaculata Blainville, Bull. Soc. Philom. Paris, 75 and 78. 

Probably Senegambia, West Africa. 

1825. Antilope suturosa Otto, Nova Acta Leop. Carol. /-'.• 519. No locality. 

1826. Antilope mytilopes H. Smith, Griffith's Cuvier Anim. Kingd. 4, pi. opposite 

p. 204 (text, 1827). Substitute for nasomaculata. 



1828. Antilope gibbosa Savi, Mem. Sci. Pisa, /; 17. Egypt. 

(Second reference: Savi, 1832, in Isis (Oken), 500. Sherborn gives Savi, 1828, 
N. Giorn. Lett. (Pisa), 16, 38: 89 and 105, but so far as the B.M. copy of 
this journal is concerned this is an error.) 

[Addax nasomaculatus addax Cretzschmar, 1826, from Dongola, Anglo-Egyptian 
Sudan, extralimital to this list, is untenable. "It may be put on record here 
that both male and female Addax imported from Dongola to the Giza 
Zoological Gardens grew fine thick winter coats" (Flower, 1932, P.Z.S. 
441) )• 

Genus ORYX Blainville, 1816 

1816. Orj'.v Blainville, Bull. Soc. Philom. Paris, ']^. Antilope oryx VaWiLi — Capra gazella 

Linnaeus (the South African Gemsbok). 
1 82 1. O^iyx Gray, London Med. Repos. i§: 307. Error for Oryx. 
19 1 8. Aegoryx Pocock, Ann. Mag. N.H. 2: 221. Cernas algazel Oken = Antilope too 

H. Smith. 

Pocock considered that Oryx tao differed so much from the other oryxes (there are 
usually held to be four species: 0. gazella, the Gemsbok; and 0. beisa,^ the Beisa 
Oryx, are African but extralimital to the Palaearctic region) that he separated it 
generically as Aegoryx. 

The grounds for this were the possession of a preorbital gland, which is said to be 
absent from the other species, a reduced rhinarium and curved horns. The preorbital 
gland is a thickening of the skin some 30 mm. long and 6 mm. thick. It is a 
superficial structure having no influence on the skull, i.e. there is no trace of a 
lachrymal pit. The difference between the rhinarium of 0. tao and 0. gazella as 
figured by Pocock seems slight, and the curved horns of tao do not seem to be 
an important character seeing that the horns in the other species show a slight 

We do not therefore consider that the differences between tao and the other species 
amount to more than specific ones. In fact, if all four species are compared it appears 
that the only one which might be considered as differing subgenerically (or generi- 
cally) is 0. leucoryx, which is a much smaller beast than the others and has a totally 
different colour pattern so far as the body and legs are concerned. We propose, how- 
ever, to leave leucoryx in the genus Oryx, and have only drawn attention to the above 
as being relevant to an assessment of Aegoryx. 

2 species in the area covered by this list: 

Oryx leucoryx, page 385 
Oryx tao, page 386 

Oryx leucoryx Pallas, 1777 Arabian Oryx 

Approximate distribution of species: Arabia, Iraq. 

* Surely 0. beisa is nothing but a north-eastern representative subspecies of 0. gazella Linnaeus. 



Oryx leucoryx Pallas, 1777 

1777. Antilope leucoryx Pallas, Spicil. Zool. 12: 17. Arabia. 

1816. Cemas oryx Oktn, Lehrb. Naturgesch. jj, 2: 734. Island in the Gulf of "Bassora". 
Unavailable, see p. 3. 

1855. -Antilope ensicornis var. asiatka Wagner, Schreb. Saugeth. Suppl. 5.- 437. Re- 
naming o( leucoryx. 

1857. Orrx beatrix Gray, P.Z.S. 157. "Shores of Persian Gulf, or of the Red Sea." 

1869. Oryx leucoryx pallani Fitzinger, S.B. Akad. Wiss. Wien, jcj, i; 178. Renaming 
of leucorvx. 

(?) 1934. Oryx leucoryx latipcs Pocock, .A.nn. Mag. N.H. /^.- 636; 1935, Ann. Mag. 
N.H. ij: 464. \\'adi Ghudun, approximately 18'^ N., 53°3o' E., Southern 
Arabia. \\"e regard this form as of doubtful validity. 

Oryx tao H. Smith, 1827 Scimitar-horned Oryx 

Approximate distribution of species: the Sahara, from .Senegal, Rio de Oro and 

Northern Nigeria to the Anglo-Egsptian Sudan, north to the Libyan Desert (Shaw, 

1933, J. Soc. Preserv. Fauna Emp., London, 20: 15). 
We do not believe this species is divisible into races. 

Oryx t.\o H. Smith, 1827 

1816. Cemas algazel Oken, L?hrb. Naturgesch. 5, 2: 741. Probably Egypt (Buffon), 

but Western Sahara according to Lydekker. Unavailable, see page 3. 
1827. Antilope tao H. Smith, Griffith's Cuvier Anim. Kingd. 4: 189. Some days' 

journey distant from the Bahr-el-Abiad, Anglo-Egyptian Sudan. 

For other extralimital synonyms, see G. Allen, 1939, Checklist African Mammals, 532. 

Subfamily A n t i 1 o p i n a e 

Genus ANTILOPE Pallas, 1766 

1766. Antilope Pallas, Misc. Zool. r. Capra cervicapra Linnaeus (fixed by Ogilby, 1836). 
1780. Cervicapra Sparrman, K. Svenska Vetensk. Akad. Handl. /.■ 281. Antilope 
cervicapra Linnaeus. 

I species: Antilope cervicapra, page 386 

Antilope cervicapra Linnaeus, 1758 Blackbuck 

Approximate distribution of species: India, from the Punjab, Kathiawar and Sind, 
eastwards to Bengal and southwards to Cape Comorin. 

Zukowsky (1927 and 1928) in a study based on over a hundred living specimens 
whose provenance was known, recognized four forms which he referred to as species 
but which are here treated as races of the same species. He distinguished these forms 
by the extent of the dark marking, greatest in hagcnbecki and least in cervicapra, by the 
degree of divergence of the horns, and by the tightness or openness of their spiral, 
combined with the number of spiral turns and the overall length of the horns. The 
largest horns and the tightest spiral occur in rajputanae and centralis, and the shortest 
with the most open spiral in cervicapra. 



Antilope cervicapra cervicapra Linnaeus, 1758 

1758. Capra cervicapra Linnaeus, Syst. Nat. loth ed. /; 69. Inland of Trivandrum, 

Travancore, India. (Zukowsky, 1927.) 
1816. Cemas strepsiceros Oken, Lehrb. Naturgesch. 5, 2: 732. India. 
1843. Cervicapra bezoartica Gray, List Mamm. B.M. 159. India. 
Range: northwards, presumably to the southern limit of the range oi centralis. 

Antilope cervicapra rupicapra Miiller, 1776 

1776. Antilope rupicapra Muller, Natursyst. Suppl. 56. Bengal. (Not preoccupied by 

Capra rupicapra Linnaeus, and supersedes hagenbecki Zukowsky.) 
1830. Antilope bilineata Gray, Illustr. Ind. Zool. /, pi. 12. India, probably Bengal. 
1927. Antilope hagenbecki Zukowsky, in Hagenbeck, Illustr. Tier. u. Menschenwelt, 2: 

125. Bengal. 
Range: nearly to Agra, in United Provinces, India. 

Antilope cervicapra rajputanae Zukowsky, 1927 

1927. Antilope rajputanae Zukowsky, in Hagenbeck, Illustr. Tier. u. Menschenwelt, 2: 

125. Neighbourhood of Bahawalpur, borders of Rajputana and Punjab. 
Range: Rajputana and Punjab. 

Antilope cervicapra centralis Zukowsky, 1928 

1928. Antilope centralis Zukowsky, in Hagenbeck, Illustr. Tier. u. Menschenwelt, j.- 

60. Gwalior, India. Range: along the southern limit of the range oi raj- 
putanae and extending an unknown distance to the south. 

Genus PROCAPRA Hodgson, 1846 

1846. Procapra Hodgson, J. Asiat. Soc. Bengal, /j.- 334. Procapra picticaudata Hodgson. 
1918. Prodorcas Pocock, Ann. Mag. N.H. 2: 130. Antilope gullurosa Pallas. Valid as a 

2 species : Procapra gutturosa, page 388 

Procapra picticaudata, page 388 

This genus contains two aberrant species formerly referred to Gazella. Lydekker, 
1914, Cat. Ungulate Mamm. B.M. j: 37, considered it was inadvisable to give Procapra 
more than subgeneric rank, but it was adopted by Pocock, and more lately by G. 
Allen and by Bobrinskii. Pocock restricted it to the type, and erected Prodorcas for the 
species P. gutturosa, which differs in glandular details from picticaudata, and seems in 
some respects to connect that species with aberrant species of Gazella like G. sub- 
gutturosa. G. Allen followed Pocock, and in his work on Mongolian and Chinese 
mammals gave Prodorcas generic rank. Simpson and Bobrinskii both seem to ignore it 
entirely. Probably subgeneric status is the most convenient treatment. Procapra differs 
from Gazella cranially very much as does Capricornis from Naemorhedus, so far as we 
have had the opportunity to examine skulls. G. Allen also gave a character of the 
nasals (p. 1209) which would separate Procapra from Gazella, and this character also 
holds fairly well when Procapra is compared with all Palaearctic species of Gazella 
available in the British Museum. 


Subgenus PROCAPRA Hodgson, 1846 

Procapra picticaudata Hodgson, 1846 Tibetan Gazelle; Goa 

Approximate distribution of species: Tibet, Southern Mongolia, Kansu and 
Szechuan in China, south to Ladak and the hills north of Kumaon and Sikkim. 
Perhaps also to Chinese Turkestan. 

Procapra picticaudata picticaudata Hodgson, 1846 

1846. Procapra picttcaudata Hodgson, J. Asiat. Soc. Bengal, /j; 334, pi. 2. Hundes 
district of Tibet. Range: Tibet and immediately adjacent parts of the Indian 
Himalayas to the south, and Szechuan in the east. 

Procapra picticaudata przewalskii Buchner, i8gi 

i8f)i. GazeUa przewalskii Buchner, Melanges Biol. St. Petersb. i^: 161. Southern 

Ordos Desert, Mongolia. 
1875. Antilopf guttiirosa Przewalski, Mongolia, /.• 18. Not of Pallas, 1777. Southern 

Ordos Desert, Mongolia. 
1888. Antilope cuvieri Przewalski, Fourth Journey in C. Asia, i 10. Not of Ogilby, 

1 84 1. Southern Ordos Desert, Mongolia. 
Range: Southern Mongolia, Northern Kansu, perhaps C'hinese Turkestan. 

Subgenus PRODORCAS Pocock, 1918 

Procapra gutturosa Pallas, 1777 Mongolian Gazelle; Zeren 

Approximate distribution of species: South-Eastern Transbaikalia, and Chuiskaya 
Steppe, South-Eastern Siberian Altai; Mongolia, Kansu, possibly (or formerly) 
Northern C'.hihli, China. 

Procapra gutturosa gutturosa Pallas, 1777 

1777. Aniilope gutturosa Pallas, Spic. Zool. 12, 46, pi. 2. Transbaikalia. (G. Allen, 
U)4o, Mamin. China & Mongolia, 121 1, fixed the type locality as the upper 
River Onon, Southern Transbaikalia.) 

1777. Antilope orientalis Erxleben, Syst. Regn. Anim. 288. Renaming oi gutturosa. 

Procapra gutturosa altaica Hollister, 1913 

1 91 3. Procapra altaica Hollister, Smith. Misc. Coll. 60, 19: i. Suok Plains, near south 
end of Bain-Chagan Pass, Little Altai, Mongolia. 

Genus GAZELLA lihiinville, 181G 

1816. Gazetlii Blainville, Bull. Soc. Philorn. Paris, 75. Capra dorcas Linnaeus. (Opinion 
108 of Liternational C^ommission on Zoological Nomenclature.) 



182 1. Dorcas Gray, London Med. Repos. /j; 307. Capra dorcas Linnaeus. 

1844. Leptoceros Wagner, Schreb. Saugeth. Suppl. 4: 422. Antilope leptoceros Cuvier. 

Not of Leach, 181 7. 
1847. T^''ogops Hodgson, J. Asiat. Soc. Bengal, 16: 695. Antilope bennettii Sykes. Not 

Tragops Wagler, 1830 (a reptile). 
1869. Tragopsis Fitzinger, S.B. Akad. Wiss. Wien, jp, i : 157. Antilope bennettii Sykes. 
i86g. Eudorcas Fitzinger, loc. cit. 159. Gazella laevipes var. a Sundevall. 
1885. Manger Lataste, Actes Soc. Linn. Bordeaux, jg: 183. Antilope (Dama)mhorr 

Bennett. Valid as a subgenus. 
1898. Korin Sclater & Thomas, Book of Antelopes, j: 65. Gazella riififrons Gray. 
1907. Matschiea Knottnerus-Meyer, Arch. Naturgesch. 75, i : 57. Gazella granti 

Brooke, from Tanganyika. 
Trachelocele^ suhgen. nov. Type species: Antilope subgutturosa Guldcnstaedt. Valid 

as a subgenus. 

Subgenus Trachelocele: Females normally hornless. Males with goitre-like 
swelling on throat during rutting season. 

Subgenus Gazella: Females with well-developed horns. No goitre-like swelling 
on throat in males. Small or medium-sized species; white of rump not 
intruding much into fawn of body. 

Subgenus Manger: Females with well developed horns. Large species; white of 
rump intruding more or less extensively into fawn of body. 

6 species in the area covered by this list : 

Gazella dama, page 394 
Gazella dorcas, page 391 
Gazella gazella, page 392 
Gazella leptoceros, page 393 
Gazella rvfifrons, page 394 
Gazella subgutturosa, page 390 

This is a difficult genus, with too many standing specific names at the present day. 
It seems that there are six valid species in the region now under discussion. These 
species are well figured in Sclater & Thomas, 1898, Book of Antelopes, j. Two of 
them, differing in colour details as indicated by Lydekker and well figured by 
Sclater & Thomas, are largely extralimital (African), but both occur in North- West 
Africa, or did until recently; these are rufifrons and dama, the latter being one of the 
species separated subgenerically as Manger. 

Of the more truly Palaearctic species, subgutturosa stands apart in that the females 
are hornless, or at most have mere rudiments of horns, whereas the females of the 
other species have distinct horns, though these are much smaller and more slenderly 
built than in the males. The species subgutturosa also differs from the other species of 
Gazella in that the male develops a goitre-like swelling on the throat in the breeding 
season. For these reasons we consider that Gazella subgutturosa should be separated 
subgenerically as indicated above. 

' From TpaxTiAos, throat; and KTi^r], tumour. Gender: feminine. 


PALAEARC;TIC and INDIAX mammals 1758-1946 

Then there arc the three common species of gazelle which range across North 
Africa and Arabia : G. gazella, G. dorcas and G. leptoceros. 

I i) G. gazella (the Arabic "Idmi"; "Mountain Gazelle" of North Africa) is a dark 
form about 70 cm. high at the shoulder, with much red-brown in its coat and 
no pure white on its face. Its habitat is chiefly in the valleys of the foothills. 
If our views are correct, that G. bennetli should probably be regarded as a 
race of G. gazella, then this gazelle, unlike the next two, ranges beyond 
Arabia into India. 

(2) G. dorcas (the Arabic '"Rhezel" or "Hamar" in North Africa and "Afri" in 

Sinai and Arabia; Dorcas Gazelle) is the smallest of these three species, 
about 54-60 cm. at the shoulder, and has a white stripe running down each 
side of the forehead and the bridge of the nose. It is paler in colour than the 
Idmi. The habitat is in the open plains. 

(3) G. leptoceros (the ,\rabic "Rhim"; Slender-horned or Lodcr's Gazelle) is lighter 

in colour than the Dorcas and the face has more white on it; the dark parts 
of the face are so pale that the gazelline face pattern is not well marked. The 
hooves are longer and narrower than in the other two species. This gazelle is 
as large as the Idmi, or larger. It is confined to the true sand areas such as 
the "ergs" of Algeria and the "nufud" of Arabia. 

The horns in gazella are smaller and stouter than in leptoceros, and those oi^ dorcas 
seem to be intermediate. But the more material accumulates the more it becomes 
apparent that the shape of the horn is unreliable as a diagnostic feature (cf Lavauden, 
iC)26, Bull. Soc. N.H. Afr. Nord. ij: 11, and Morrison-Scott, 1939, Novit. ^ool. .^i: 

Subi;cnus TRACHELOCELE Ellcrman & Morrison-Scott, 1951 

Gazella subgutturosa Guldenstaedt, 1780 Goitred Gazelle; Persian Gazelle 

Approximate distribution of species: Transcaucasia, Russian Turkestan, where 
widely distributed (absent from Fergana \''alley), northwards to Tarbagatai Moun- 
tains, eastern half of Lake Balkash, basin of the Sarui-Su, thence west to the northern 
part of Ust-Urt (Bobrinskii). Mongolia, Chinese Turkestan, Northern Tibet. Persia, 
Euphrates Valley, Afghanistan, Baluchistan. 

Gazell.\ subgutturos.'v subgutturosa Giildenstaedt, 1780 

1780. Antilope subgutturosa Guldenstaedt, Acta Ac. Sci. Petrop. i/j8, i : 251. North- 
Western Persia. 
1843. Antilope dorcas var. persica Gray, List Mamm. B.M. 160. 
1900. Gazella subgutturosa typica Lydekker, Great & Small Game India, 180. 

Range: Afghanistan, Euphrates Valley, Persia and Russian Turkestan. 


Gazella subgutturosa yarkandensis Blanford, 1875. Saikik Gazelle 

1875. Gazella subgutturosa \'z.r. yarkandensis Blanford, J. Asiat. Soc. Bengal, 44, 2: 112. 

Plains of Yarkand, Chinese Turkestan. 
1910. Gazella yarcandensis Lydekker, Nature, 8^: 202. 

Gazella subgutturosa hillieriana Heude, 1894 

1894. Gazella hillieriana Heude, Mem. H.N. Emp. Chin. 2: 245, pi. 36. ? Gobi Desert, 

Eastern Mongolia. 
1894. Gazella mongolica Heude, loc. cit. pi. 37. 

Range: Mongolia. Synonyms of the typical race according to Lydekker; G. Allen 
(1940) revives the name for the Mongolian form. 

Gazella subgutturosa sairensis_ Lydekker, 1900 

1900. Gazella subgutturosa sairensis Lydekker, Great & Small Game of India, 184. 
Saiar Mountains, Zungaria. 

Gazella subgutturosa seistanica Lydekker, 19 10 

1910. Gazella seistanica Lydekker, Nature, 8j: 202. Seistan, Eastern Persia. Range: 
to Baluchistan. 

Gazella subgutturosa reginae Adlerberg, 1931 

1931. Gazella subgutturosa reginae Adlerberg, C.R. Acad. Sci. U.R.S.S. 327. North- 
Western Tsaidam, Northern Tibet. 

Subgenus GAZELLA Blainville, 181 6 

Gazella dorcas Linnaeus, 1758 Dorcas Gazelle 

"Rhezel" (North Africa); "Afri" (Sinai and Arabia) 

Approximate distribution of species: Northern Africa, from Rio de Oro, Morocco, 
Algeria, Tunisia, Libya, Egypt, east to Sinai, Palestine, Syria, Arabia, and south to 
the Sudan, Abyssinia, Lake Chad region. 

Gazella dorcas dorcas Linnaeus, 1758 

1758. Capra dorcas Linnaeus, Syst. Nat. loth ed. /.• 69. Lower Egypt (Blaine, 1913, 

Ann. Mag. N.H. //.• 292). 
1766. Antilope kevella Pallas, Misc. Zool. 7. Based on "Le Kevel" of Buffon, 1764. 
1766. Antilope corinna Pallas, loc. cit. Based on "La Corinne" of Buffon, 1764. 
1816. Cemas maculata Oken, Lehrb. Naturgesch. 5.- 738. Senegal. 
1869. Gazella dorcas sundevalli Fitzinger, S.B. Akad. VViss. Wien, ^g, i: 159. North 

Africa. (Naming of G. dorcas var. y Sundevall.) 
Range : Rio de Oro to Egypt and the Sudan. 

Gazella dorcas neglecta Lavauden, 1926 

1926. Gazella dorcas neglecta Lavauden, Bull. Soc. H.N. Afrique du Nord, ly: 16. 
Text figs. 2, 3 and 4. Plateau de Tadmeit, Central Algerian Sahara. 



Gazella dori:as massaesyla Cabrera, 1928 

1928. Gazella dorcas massaesyla Cabrera, J. Mammal, (j: 242. High plateaux of the Rif, 

Moroecci, sDuth of Spanish-French frontier. 

1929. Gazella dorcas eahrerai ]o\eii\iA, Bull. See. Zool. France, §4: 440. Substitute for 

massaesyla thought to be preoccupied by massaesilia Pomel, 1894, Carte Geol. 
de rAlgcrie, Lcs Antilopes, 21. 

G.'^zell.'l DORCAS SAUDiY.x Carruthcrs & Schwarz, 1935 

1935. Gazella gazella saudiya Carruthers & Schwarz, P.Z.S. 155. Dhalm, about 150 
miles north-east of Mecca, Arabia. Range: Sinai, Arabia, Palestine. 

Gazella gazella Pallas, 1766 Mountain Gazelle (North Africa) 

"Idmi" (North Africa and Arabia); Chinkara (India) 

Approximate distribution of species: Rio de Oro, Morocco, Algeria, Western 
Tunis; Sinai and Arabia, south to Aden, Eastern Persia, Palestine, Syria; Baluchistan, 
Punjab, Sind, Nepal, United Provinces, Rajputana, Cutch, Kathiawar, to a little 
south of the Krishna (Kistna) River (which forms the border between Hyderabad 
and Madras), India. 

Gazella gazella gazella Pallas, 1766 

1766. Antilope gazella Pallas, Misc. Zool. 7. Syria. 

1904. Gazella mm//// Thomas, Abstr. P.Z.S. No. 12, 19; 1905, P.Z.S. igo4, 2: 347. 

Hizmeh, north of Jerusalem, Palestine. 
Range : Syria and Palestine. 

Gazella gazella arabica Lichtenstcin, 1827 

1827. Antilope arabica Lichtenstein, Darstellung Saugeth. pi. 6. Farsan Island, on 
Arabian coast of Red Sea. (Some authors have "Sinai", on the ground that 
Lichtenstein's description was based on Ehrenberg's manuscript, and 
Ehrenberg's plate, published in 1828, is marked "Sinai".) 

1827. Antilope cora H. Smith, Griffith's Cuvier Anim. Kingd. 4: 216. Persian Gulf. 

1874. Gazella miiscalensis Brooke, P.Z.S. 142, pi. 22. Muscat, Oman, Eastern Arabia. 

1906. Gazella arabica crlangeri Neumann, S.B. Ges. Naturf Fr. Berlin, 244. Lahej, 
near Aden, South-\Vestern Arabia. 

1906. Gazella arabica rueppelli Neumann, loc. cit. Sinai Peninsula. 

iqio. Gazella arabica Irpica \Vard, Rec. Big Game, ed. 6, 251. 

1927. Gazella arabica hanishi Dollman, Abstr. P.Z.S. No. 291 : i; 1928, P.Z.S. igsy: 
1005. Great Hanish Island, Red Sea. 

Gazella gazella bennetti Sykes, 1831 

1831. Antilope beruiettii Sykea, P.Z.S. 1830-31: 104. Deccan, India. 

1839. Antilope arabica Elliot, Madras J. Lit. 10: 223. Not of Lichtenstein, 1827. 

1842. Gazella christii Blyth, J. Asiat. Soc. Bengal, //.• 452. Thar (or Indian Desert), 


1843. Antilope hazenna I. Gcoffroy, in Jacciucmont, Voy. Indc, 4: 74. Malwa, Central 

(?) 1873. Gazella fuscifrons Blanford, P.Z.S. 317. Jalk, on fringe of Seistan Desert, 
Eastern Persia. 



1908. Gazella yarkandensis kennioni Lydekker, Field, ///; 499. Kain, Afghan frontier 
of Persia. 

191 1. Gazella hayi Lydekker, P.Z.S. 961. "As the result of an unfortunate accident, 
namely the transposition of the registration labels of two gazelles received 
simultaneously at the British Museum, I find that I have described and 
figured a specimen of the Seistan Gazella fuscifrons as a new African species 
under the name G. hayi" (Lydekker, 1912, P.Z.S. 911). 

Range: Eastern Persia, and Indian range of the species above. 

Gazella gazella cuvieri Ogilby, 1841 

1841. Antilope cuvieri Ogilby, P.Z.S. 1840: 35. Mogador, Morocco. 

1804. Gazella comma Lacepede & Cuvier, Menag. Mus. H.N. Paris, plate and text. 

Not of Pallas, 1766. Constantine, Algeria. 
1850. Gazella vera Gray, Gleanings Menag. Knowsley Hall, pi. 3. 
1853. Gazella cineraceus Temminck, Esquis. Zool. sur la Cote de Guine, 193. No 

i860. Gazella kevella Tristram, The Great Sahara, 387. Not of Pallas, 1766. Atlas 

Mountains, south of Teniet el Haad, West of Algeria. 

Range: Mountains of Morocco, Algeria, Tunis. 

Gazella leptoceros F. Cuvier, 1842 Slender-horned Gazelle; Loder's Gazelle 

"Rhim" (North Africa and Arabia); "Gazal abiad" (Egypt) 

Approximate distribution of species: Algeria, Libya, Egypt, the Sudan and Arabia. 

Gazella leptoceros leptoceros F. Cuvier, 1842 

1842. Antilope leptoceros Cuvier, in Geoffroy & Cuvier, Hist. Nat. Mamm. 4: lixraison 

72. Antilope a longues comes, 2, and pis. 373, 374. "Sennaar" (probably the 
desert between Giza and Wadi Natron, Lower Egypt, as the type specimen 
was brought to Paris by James Burton, circa 1833. Flower, 1932). 

1869. Leptoceros abu harab Fitzinger, S.B. Akad. Wiss. Wien, §g, i: 160. Libyan 

1869. Leptoceros cuvieri Fitzinger, loc. cit. Renaming oi leptoceros. Nee Ogilby, 1841. 

(?) 1894. Gazella loderi Thomas, Ann. Mag. N.H. i^: 452. Sand dunes of Oued Souf, 
100 miles south of Biskra, Northern Algeria. Often regarded as a race of 
leptoceros, but "points of difference from typical race not known" (Lydekker). 

1898. Gazella leptoceros typica Sclater & Thomas, Book of Antelopes, 5.- 149. 

Gazella leptoceros marica Thomas, 1897 

1897. Gazella marica Thomas, Ann. Mag. N.H. ig: 162. Nejd, Central Arabia- 
Range : sand areas of Arabia. 

Thomas said that marica agreed with leptoceros in the almost complete absence of 
gazelline face markings and general pale colour, but that it was smaller and had 
shorter and differently shaped horns. But Thomas had only four specimens and the 
only adult male had the horns sawn off. Other reasons for regarding marica as a race 
oileptoceros is that its habitat is restricted to sandy dunes in the same way as the latter, 
and its Arabic name is the same — "Rhim". Thomas's chief point of difference, that of 



the small size of marica, is belied by the fact that the Arabs of Arabia regard "Rhim" 
as the largest of their three gazelles. 

Gazella rufifrons Gray, 1846 Korin or Red-fronted Gazelle 

Approximate distribution of species: Senegal, Gambia, Northern Nigeria, region 
of Lake Chad, the Sudan. Possibly, not certainly, still existing in Algeria. Heim de 
Balsac, 193(3, Bull. Biol. France et Belgique, Suppl. 21: 88, regards rufina, of which only 
three or four specimens are known, as a species distinct from rufifrons. 

(Gazella riififrons rufifrons Gray, 1846. Extralimital) 
1846. Gazella rufifrons Gray, Ann. Mag. N.H. 18: 214. Senegal. 


1894. Gazella rufina Thomas, P.Z.S. 467. "Probably the interior of Algeria"; type 

purchased at Algiers. Now believed to be extinct, but there may be a herd 
in the Chelif district, between Oran and Algiers (Heim de Balsac, 1936). 

Gazella (?) rufifrons pallaryi Pomcl, 1895 

1895. Antilope (Dorcas) pallaryi Pomel, Paleontologie-Les Antilopes Pallas, 9. Type 

and only specimen obtained from a merchant in Oran, Algeria. 

Subgenus NANGER Latastc, 1885 

Of the three species referred by Lydekker and others to this subgenus, only one, 
the earliest named, enters the present region. 

Gazella dama Pallas, 1 766 Dama Gazelle 

Approximate distribution of species: Senegal, Lake Chad district, the Sudan, 
northwards to Morocco and Rio de Oro. 

(Gazella d.\ma dama Pallas, 1766. Extralimital) 

I 766. Antilope dama Pallas, Misc. Zool. 5. Probably the vicinity of Lake Chad, French 
Equatorial Africa.' 

Gazella dama mhorr Bennett, 1833 

1833. Antilope {Dama) mhorr Bennett, P.Z.S. 2. Wednun, near Tafilat, Mogador, 

1846. Gazella moiir Gray, Ann. ^L'^g. N.H. 18: 231. Emendation oi mhnrr. 

Gazella dama lozanoi Morales Agacino, 1934 

1934. Gazella dama lozanoi Morales Agacino, Bol. Soc. Esp. H.N. 34: 454, pi. 35, fig. i. 
Villa Cisneros, Rio de Oro. 

' 1847. Aniilofie dama var. occtdenlalis Sundevall, K. Svenska Vetensk. .'\kad. Hand). 1845: 266. 
Currently listed as a synonym ol' G. d. dama. Sundevall gives "var. ocadentalis" twice under Antilope 
dama. once with locality "Senaar, Egyptus", and a second time with "Senegal, Marocco". It may 
w.-U be that the first "var. occidentalis" is a misprint lor "orienlalis", but he does not say so in the list of 
cfirrigenda given on p. 324. 



Subfamily C a p r i n a e 

As understood by Simpson (1945) this contains four tribes, typified by Saiga (with 
Pantholops) ; Budorcas (allied to the Nearctic Ovibos in Simpson's list); Rupicapra, with 
immediate allies; and Capra, with immediate allies. These four groups are usually 
given subfamily rank. Sometimes Saiga and Pantholops are considered to be more 
closely allied to the Antilopinae. The very remarkable structure of the skull in the 
region of the nasal aperture in Saiga is well figured in Bobrinskii (1944). 

Genus PANTHOLOPS Hodgson, 1834 
1834. Pantholops Hodgson, P.Z.S. 81. Antelope hodgsonii Abel. 
I species : Pantholops hodgsoni, page 395 

Pantholops hodgsoni Abel, 1826 Chiru; Tibetan Antelope 

Approximate distribution of species: Tibet; Ladak. "The only spot in Indian 
territory in which (Chiru) are found is the Chang Chen Mo Valley (Northern Ladak) 
into which they cross from Tibet by way of the Lanak La Pass, at the head of the 
valley" (Prater). 

Pantholops hodgsoni Abel, 1826 

1826. Antelope he dgsonii Abel, Calcutta Govt. Gazette, 1826. {J^.V.}: Phil. Mag. 68: 

234. Tingri Maidan, Arrun Valley, Kooti Pass, Tibet. 

1827. Antilope kemas H. Smith, Griffith's Cuvier Anim. Kingd. ./.• 196. Central Asia. 
1827, Antilope chiru Lesson, Man. Mamm. 371. Nepal. 

Genus SAIGA Gray, 1843 

1843. Saiga Gray, List Mamm. B.M., xxvi. Capra tatarica Linnaeus. 

1843. Siaga Gray, List Mamm. B.M. 160. 

1844. Colus Wagner, Schreb. Saugeth. Suppl. ^: 419. Antilope saiga Pallas = Capra 

tatarica Linnaeus. 

I species: Saiga tatarica, page 395 

Saiga tatarica Linnaeus, 1 766 Saiga 

Approximate distribution of species: "Nowadays the saiga only remains in the area 
stretching from the right-bank steppe of the Lower Volga across Kazakstan to Zun- 
garia, inclusive. Even in that area however its distribution within the U.S.S.R. is 
not continuous, but divided into separate districts: (i) Kalmuikia; (2) the Volga- 
Ural steppes; (3) the steppes between the River Emba and lower River Ural; 


1'ai,aearc:tk: and lxdian mammals 1758-194G 

(4) Biizachi Peninsula; (5) Northern Ust-Urt ? (it apparently fails to occur in the 
more southern parts of the Ust-Urt and only in the winter occasionally visits the 
Kara-bou,e;az area) ; (6) east of the Amu-Darya delta; (7) Barsa-Kalmes Island, in the 
Sea of Aral; (8) the lower Syr-Darya area, Karsakpai area, the lower and middle 
Sarui-Su and the Godolnara steppe (Betpakdala) ; (9) the steppes between Lake 
Balkash and the Rivers Hi and Karatal; (10) the Ala-Kul basin; (11) the north of 
Zaisan basin. Rare everywhere in the U.S.S.R. Hunting rif it everywhere forbidden" 
(Bobrinskii, 1944). In addition, a form has recently been named from Mongolia. This 
differs from ,S'. tatarka in smaller size, detailed structure of horns, and some cranial 
details. From descriptions it might equally well be regarded as a species or as a very 
distinct race of tatarka. 

The name Ihex imberbis Gmclin, 1760, Nov. Comment. Acad. Sci. Fctrop. ^: 345 (and 
1761, y: Summarium, 39), Tara, on Irtish River, Siberia, has been used for the 
Saiga but is unavailable, since in this particular work Gmelin is not consistently 

Saig.-\ t.marica tatarica Linnaeus, 1 766 

1760. Ihix imberbis Gmelin, Nov. Comment. Acad. Sci. Petrnp. j.- 345 (and 1 761, y: 

Summarium, 39). Tara, on River Irtish, Siberia. (Unavailable.) 
1766. Ca/ira tatariea Linnaeus, Syst. Nat. 12th ed. /.• 97. Ural Steppes, Russia. 

1766. Antilope saiga Pallas, Misc. Zool. 6. Renaming o( Ibex imberbis Gmelin. 

1767. Antilope scvlhiea Pallas, Spic. Zool. /.• 9. Renaming o^ imberbis. 

1768. Capra sajga Forster, Philos. Trans. ;jy: 344. Vr)Iga Basin, Russia. 

1816. Cemas coins Oken, Lehrb. Naturgcsch. jj, 2: 736. Renaming oH Ibex imberbis. 
Range: Russian and Siberian range of species. 

Saiga ( .■') t.\tarica mongolica Bannikov, 1946 

1946. Saiga mongolica Bannikov, C.R. Acad. Sci. U.R.S.S. 5/.- 401. One hundred and 
fifty kilometres west of Bayan Somon, Shargin Gobi, Dukhmen-tala, 
Western Mongolia. 

Genus BUDORCAS Hodgson, 1850 
1850. fliidoreas Hodgson, J. A=iat. Soc. Bengal, ir/: 65. Budnreas taxieolor Hodgson. 
I spc( ies: Budorcas taxieolor, page 39B 

Budorcas taxieolor Hodgson, 1850 Takin 

.\l)l)r(iximate distribution of species: Mishnii, Bhutan, and possibly Northern 
I'jurma; st.ites of Szcchuan and Shensi (possibly \'vninan on Burma border) and 
probably iiitu .S